





























































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































































Class HQ 750 
Book.___3_ Xii 

CojpghtN?_ ISjLiQs 


COEBRIGHT DEPOSIT- 















































. 


















« 








































































































































































































































































* 
































'V 
































































































































































































Eugenics, 

Genetics and the Family 

VOLUME I 


SCIENTIFIC PAPERS OF THE 


r\jLur __ 


JLaa^cvU-tvcco-- 

SECOND INTERNATIONAL CONGRESS OF EUGENICS 

A 


HELD AT 

AMERICAN MUSEUM OF NATURAL HISTORY, NEW YORK 
September 22-28, 1921 


Committee on Publication 
Charles B. Davenport, Chairman 
Henry Fairfield Osborn, Ex Officio 
Clark Wissler 
H. H. Laughlin, Secretary 


> ) 


BALTIMORE 

WILLIAMS & WILKINS COMPANY 
1923 


HQ iso 
■AzTu 

/*? & / CXs 


/ 


Copyright 1923 

WILLIAMS & WILKINS COMPANY 


Mode in United States of America 

All rights reserved, including that of translation 
into foreign languages, including the 
Scandinavian 



COMPOSED AND PRINTED AT THE 


WAVERLY PRESS 

Bv the Williams & Wilkins Company 

BALTIMORE, MARYLAND. U. S. A. 


MAH'I’23 

©C1AG<JS4C3 


a . 

































































■ 








































Alexander Graham Bell 


Honorary president of the Second International Congress of Eugenics, and pioneer 
investigator in the field of human heredity. 




CONTENTS 


Preface. ix 

Scientific Papers: 

1. Address of Welcome, Dr. Henry Fairfield Osborn. American Museum of 

Natural History. 1 

2. Aims and Methods of Eugenical Societies, Major Leonard Darwin. Pres. 

Eugenics Education Society, England. 5 

3. Research in Eugenics, Dr. Charles Benedict Davenport. Carnegie Institu¬ 

tion of Washington, Cold Spring Harbor, N. Y. 20 

4. Genetique et Adaptation, Dr. L. Cu6not. Professor of Zoology, University 

of Nancy, Nancy, France. 29 

5. Inheritance in Unicellular Organisms, Dr. H. S. Jennings. Johns Hopkins 

University. 59 

6. Evolution of the Chromosome Complex, Dr. C. E. McClung. University 

of Pennsylvania. 65 

7. Aberrations in Chromosomal Materials, Dr. Calvin B. Bridges. Columbia 

University, New York City. 76 

8. Variations in the Jimson Weed (Datura Stramonium), Dr. A. F. Blakeslee. 

Carnegie Institution of Washington, Cold Spring Harbor, L. I., N. Y. 82 

9. The Attraction between Homologous Chromosomes, Mr. John Belling. 

Carnegie Institution of Washington, Cold Spring Harbor, L. I., N. Y. 84 

10. Linkage with Lethal Factors in the Solution of the Oenothera Problem, 

Dr. George H. Shull. Princeton University. 86 

11. A New Type of Variability in Plants, Dr. R. Ruggles Gates, University of 

London. 100 

12. The Analysis of Genetic Differences through Haploid Parthenogenesis, 

Dr. P. W. Whiting. Child Welfare Research Station, State Univ. 
of Iowa. 102 

13. Mutation, Dr. H. J. Muller. University of Texas, Austin, Texas. 106 

14. Reverse Mutation, Dr. Charles Zeleny. University of Illinois. 113 

15. Darwinian Evolution by Mutation, R. A. Fisher, M. A., Rothemsted Experi¬ 

mental Station, Harpenden, England. 115 

16. The Frequency of Mutation and the Incidence of Hereditary Traits in Man, 

Dr. C. H. Danforth. Washington University Medical School, St. 
Louis. 120 

17. A Simple Explanation of the Hereditary Mechanism, Dr. Louis Legrand, 

Tunis, Africa. 129 

18. Sex Determination in Rotifers, Dr. A. Franklin Shull. University of 

Michigan, Ann Arbor, Mich. 138 

19. Some Data on Control of Sex in Cladocera, Dr. Arthur M. Banta and Mr. L. 

A. Brown. Carnegie Institution of Washington, Cold Spring Harbor . 142 



















CONTENTS 


20. Disturbances in Mammalian Development Produced by Radium Emanation, 

Dr. Halsey J. Bagg. Huntington Fund for Cancer Research and Cor¬ 
nell University Medical College, N. Y. C. 150 

21. The Genetic Significance of the Alcoholic Treatment of White Rats, Dr. E. 

Carleton MacDowell. Carnegie Institution of Washington. 168 

22. Inheritance in Mental Disorders, Henry A. Cotton, M.D. N. J. State 

Hospital, Trenton. 170 

23. Comparison of the Incidence of the Supracondyloid Process in Groups with 

Normal and Abnormal Mentality, Dr. R. J. Terry and Dr. Lee D. Cady. 
Washington University School of Medicine, St. Louis. 174 

24. The Relation of Heredity to Tuberculosis, Dr. Paul A. Lewis, Phila¬ 

delphia, Pa. 178 

25. The Inheritance of Cancer in Mice, Dr. Leo Loeb. Washington University, 

St. Louis.. 182 

26. The Inheritance of a Predisposition to Cancer in Man, Dr. C. C. Little. 

Carnegie Institution of Washington, Cold Spring Harbor, L. I., N. Y.... 186 

27. Inheritance of Eye Defects, Dr. Lucien Howe. Buffalo, N. Y. 191 

28. New Data on the Genesis of Twins, R. A. Fisher, M. A. Rothemsted Experi¬ 

mental Station, Harpenden. 195 

29. Main Results of a Statistical Investigation of “Finger Prints” from 24518 

Individuals, Kristine Bonnevie. Univ. of Christiania, Norway.*.. 198 

30. The Genetics of Fecundity in the Domestic Hen, Major C. C. Hurst, Bur¬ 

bage, Leicestershire. 212 

31. Inheritance of Mental Disease, Dr. A. Meyerson, Boston, Mass. 218 

32. Inheritance of Mental Disorders, Dr. Aaron J. Rosanoff. Kings Park State 

Hospital, N.Y. 226 

33. Individual and Racial Inheritance of Musical Traits, Dr. Carl E. Seashore. 

State University of Iowa. 231 

34. An Experimental Investigation of Musical Inheritance, Hazel M. Stanton. 

State University of Iowa. 239 

35. The Consequences of War and the Birth Rate in France, Dr. M. Lucien 

March, Paris, France. 243 

36. The Effects of Inbreeding on Guinea-Pigs, Dr. Sewall Wright. Bureau of 

Animal Industry, U. S. Dept, of Agriculture, Washington, D. C. 266 

37. Is Inbreeding Injurious? Dr. Helen Dean King. Wistar Institute, 

Philadelphia. 270 

38. Intermarriage of Blood Relatives in Three Old New England Communities, 

Mrs. Ruth Moxcey Martin, Woodbury, Conn. 278 

39. Formal Inbreeding in Human Society with South American Examples, 

Dr. Herbert J. Spinden. Peabody Museum of Harvard University ... 285 

40. The Effect on the Germ Plasm of Isolation in a Mountain Section, Prof. W. 

S. Anderson. University of Kentucky. 297 

41. Developpement compare des produits successifs d’un meme couple, 

M. Etienne Rabaud. 303 

42. The Ideal Family History, Dr. Howard J. Banker, Eugenics Record Office, 

L.I.,N.Y. 306 

43. The Conification of Social Groups. Evidence from New England Families, 

Dr. Frederick Adams Woods, Massachusetts Institute of Technology. . 312 























CONTENTS 


V 


44. The Mayflower Pilgrims, Sarah Louise Kimball. California Geneological 

Society, San Francisco. 329 

45. The Learned Blacksmith—An Aristogenic Type, Dr. Howard J. Banker. 

Eugenics Record Office, Cold Spring Harbor, N. Y. 340 

46. Breve Resena Genealogica de la Familia Izquierdo, Dr. J. Joaquin Izquierdo. 

M. S. A., Mexico City, Mexico. 348 

47. The Oneida Community Experiment in Stirpiculture, Dr. Hilda Herrick 

Noyes and George Wallingford Noyes, Kenwood, N. Y. 374 

48. Some Families as Factors in Anti-Social Conditions, Dr. Amos W. Butler, 

Secretary, Board of Charities, State of Indiana, Indianapolis, Ind.387 

49. Pedigrees of Pauper Stocks, Mr. E. L. Lidbetter, London, England. 391 

50. The Tribe of Ishmael, Dr. Arthur H. Estabrook. Eugenics Record Office, 

Cold Spring Harbor, N. Y. 398 

51. Heritable Factors in Human Fitness and Their Social Control, Dr. Wilhel- 

mine E. Key. Race Betterment Foundation, Battle Creek, Mich.405 

52. A Study of 150 Adolescent Runaways, Miss Elizabeth Greene. Waverly 

House, N. Y. C,. 413 

53. Mate Selection, Dr. Roswell H. Johnson. University of Pittsburgh.416 

Discussions of papers read at the Congress. 426 















« 


» 


✓ 


\ 





I 










% 














PLATES: VOLUME I 


I. Alexander Graham Bell. Frontispiece, opposite iii 

II. Major Leonard Darwin. Frontispiece, opposite p. 5 


Plates at Back of Volume 

1. The chromosomes of man. 

2. The Catlin mark. 

3. Inheritance of order of succession in development of the carpal bones. 

4. Heredity of longevity. 

5. Pedigrees of dramatic and musical talent. 

6. Heredity of musical ability. 

7. Heredity of harelip and cleft palate. 

8. Heredity in epilepsy. 

9. Inheritance of specific iso-agglutinins. 

10. Color inheritance in corn. 

11. Measurement of physical traits. 

12. Measurement of physical and mental traits. 

13. Measurement of mental traits at Vineland. 

14. Palm and sole prints. 

15. Composite portraiture. 

16. Pedigree of John Burroughs. 

17. Pedigree of the Caesars. 

18. The Old Americans and the Tribe of Ishmael. 

19. The Jukes. 

20. The Nams. 

21. Mentality and delinquency. 

22. The brains of criminals. 

23. Marriage and birth rate in relation to immigration. 

24. Approaching extinction of Mayflower descendants. 


i 


vii 




PREFACE 


This volume contains a series of addresses and scientific papers given at 
the Second International Congress of Eugenics held in New York in Sep¬ 
tember, 1921. The three addresses speak for themselves, and are intended 
to make a popular appeal. The scientific papers are in the field of heredity 
and family studies. It is distinctive of eugenics that it is an applied 
human science, in this respect like medicine; but one dealing with the 
larger biologico-social aspects of the human race. To-day, as never before, 
we realize that at the bottom of real social progress lies the germ plasm— 
the chromosome complex which, during all these eons, has been controlling 
the progress of the organic world; created the precursor of man and, all 
hidden and unknown, has directed human evolution with a certainty that 
has made all human endeavors in this direction almost negligible. It speaks 
for the breadth of geneticists that, while recognizing that there is much said 
and done in eugenics that is not scientific, they were glad to contribute 
their special knowledge to a subject of such importance for humanity. 
Through this contribution which they have made, the methods and some 
of the results of genetical research will become better known to those who 
are applying eugenical principles. Pages 29 to 242 contain the results of 
some of the most advanced and best known investigators in genetics. They 
give an insight into the phenomenal progress that has been made in the past 
twenty years in the field of heredity. 

The particular field of human genetics has methods of its own, just be¬ 
cause human matings are not under the control of the geneticist. One of 
these distinctive methods is, accordingly, the collection of family pedigrees, 
to learn what sorts of matings have produced particular kinds of progeny, 
and, conversely, what kind of progeny have resulted from particular 
matings, bringing together particular kinds of gametes. So the analysis 
of human families is of prime importance in eugenics, and the present 
volume contains, pp. 243-390, reports on biological researches on families 
and the results of human matings. Here are discussed data on differential 
fecundity, on inbreeding, on special strains, on particular families, on experi¬ 
ments in the control of human matings, on family traits in relation to social 
conditions. 

For the publication of this volume, there is first of all to be thanked the 
contributors to it, who have given it its great intrinsic value, and who have 


IX 


X 


PREFACE 


cheerfully responded to the requests of the Publication Committee. But 
these efforts and good will alone would not have sufficed, since no publisher 
could be found who would undertake the manufacture of the book without 
financial guarantees. After some delay, these guarantees were secured and 
we are here expressing the thanks that all must feel to the special contribu¬ 
tors to the publication fund: Dr. Frederick Adams Woods, Mrs. E. H. 
Harriman, Mr. Alexander B. Coxe, Mr. Charles W. Gould, and Mr. Madison 
Grant. Thanks are due also to the Messrs. Williams & Wilkins Company 
who have assumed no little risk in undertaking the manufacture of these 
volumes, and who have spared no pains to publish the work promptly and 
in attractive form. Finally mention must be made of the painstaking 
work of Miss Luella H. Smith who prepared the index to the volumes. 


ADDRESS OF WELCOME 

HENRY FAIRFIELD OSBORN 
American Museum of Natural History 

I doubt if there has ever been a moment in the world’s history when an 
international conference on race character and betterment has been more 
important than the present. Europe, in patriotic self-sacrifice on both 
sides of the World War, has lost much of the heritage of centuries of civiliza¬ 
tion which never can be regained. In certain parts of Europe the worst 
elements of society have gained the ascendancy and threaten the destruc¬ 
tion of the best. At this moment we welcome the sound and highly trained 
judgment of Major Leonard Drawin, leader of the eugenics movement in 
Greater Britain; of Dr. Lucien March, the leading statistical authority of 
France, also leader in the eugenics movement and senior representative of 
the eugenics movement there; of Dr. Lucien Cuenot, foremost student of 
the science of heredity in France; of Dr. G. V. de Lapouge of France, the 
leading authority on racial anthropology and earnest exponent of practical 
eugenic measures by the government. Dr. Jon Alfred Mj0en of Norway is 
the leader in the vigorous movement of race hygiene in Scandinavia. 
Contributions are welcomed from other representatives of Great Britain, 
of France, of Italy, of Belgium, of the new Republic of Czecho-Slovakia, of 
our sister Republic of Cuba, and of South and Central America. The 
leading students of heredity, of statistics, of anthropology, and of eugenics 
in the United States are here to welcome their confreres from abroad. 

To each of the countries of the world, racial betterment presents a 
different aspect. To the five countries most closely engaged in the recent 
fratricidal conflict the financial and economic losses of which we hear so 
much are as nothing compared with the spiritual, intellectual, and moral 
losses which each has sustained. In the Scandinavian countries, which 
kept out of the conflict, and to a large extent in the United States the case 
is different. In Scandinavia, which I have recently visited, it is largely 
through the active efforts of leaders like Mj0en and Lundborg that there is 
a new appreciation of the spiritual, intellectual, moral, and physical value 
of the Nordic race, and that a warning is being given that it must not be too 
severely depleted by emigration. Nearly half that race is now in the 
United States. 


1 


2 


HENRY FAIRFIELD OSBORN 


In the United States we are slowly waking to the consciousness that 
education and environment do not fundamentally alter racial values. 
We are engaged in a serious struggle to maintain our historic republican 
institutions through barring the entrance of those who are unfit to share the 
duties and responsibilities of our well-founded government. The true 
spirit of American democracy that all men are born with equal rights and 
duties has been confused with the political sophistry that all men are born 
with equal character and ability to govern themselves and others , and with the 
educational sophistry that education and environment will offset the handi¬ 
cap of heredity. South America is examining into the relative value of the 
pure Spanish and Portuguese and of various degrees of racial mixture of 
Indian and Negroid blood in relation to the preservation of their republican 
institutions. 

In my recent tour through Belgium and all parts of France, I was deeply 
impressed with the very slight convergence produced by 12,000 years of 
similar environment and a thousand years of similar education upon the 
three divergent races of which France is composed,—the Mediterranean, 
the Alpine, and the Nordic. 

The constructive spirit of this Congress is to discover the virtues and 
the values of each of these minor divisions of the human species, as well as 
the needs of the major divisions, known as the Caucasian, the Mongolian, 
and the Negroid. The reason that these races are so stable and maintain 
their original character so stoutly is that the most stable form of matter 
which has thus far been discovered is the germ plasm on which heredity 
depends. This outstanding fact of heredity will be brought out in the 
First Section of the Congress. As a palaeontologist and geologist, as well 
as something of a biologist, I find no form of matter so stable in nature as 
that on which heredity depends—consequently the selection, preservation, 
and multiplication of the best heredity is a patriotic duty of first importance. 
In the selection of the best we should know no prejudice. If we extenuate 
nothing, we write down nothing in malice. The 500,000 years of human 
evolution, under widely different environmental conditions, have impressed 
certain distinctive virtues as well as faults on each race. In the matter of 
racial virtues, my opinion is that from biological principles there is little 
promise in the “melting pot” theory. Put three races together, you are as 
likely to unite the vices of all three as the virtues. This opinion, however, 
awaits the experimental proof or disproof which will be presented by re¬ 
searches such as those of Doctor Sullivan in the Hawaiian Islands. For 
the world’s work, give me a pure-blooded Negro, a pure-blooded Mongol, 
a pure-blooded Slav, a pure-blooded Nordic, and ascertain through observa- 


ADDRESS OF WELCOME 


3 


tion and experiment what each race is best fitted to accomplish in the 
world’s economy. If the Negro fails in government, he may become a 
fine agriculturist or a fine mechanic. The Chinese and the Japanese have 
demonstrated in the history of their respective countries a range of ability 
in art, literature, and industry quite equal to our own in certain arts, and 
greatly superior to our own in other arts, like ceramics. Let each race con¬ 
sider its own problems and demonstrate its own fitness. 

Our Fourth Section is devoted to the state. The right of the state to 
safeguard the character and integrity of the race or races on which its 
future depends is, to my mind, as incontestable as the right of the state to 
safeguard the health and morals of its people. As science has enlightened 
government in the prevention and spread of disease, it must also enlighten 
government in the prevention of the spread and multiplication of worthless 
members of society, the spread of feeble-mindedness, of idiocy, and of all 
moral and intellectual as well as physical diseases. 

I would not anticipate the findings of any of the four sections into which 
the work of the Congress is divided, but I would express my opinion that 
the monogamous family, i.e., one husband, one wife, is to be maintained and 
safeguarded by the state as well as by religion as a natural and hence as a 
patriotic institution. In Doctor Lowie’s very able recent work, “ Primitive 
Society,” it is shown that in general the family is safeguarded; that the 
natural instinct so widely prevalent among all social lower orders of animals 
to preserve the family at all costs dominates the elementary morals of 
primitive races. It is not an exaggeration to say that many tendencies in 
recent social development, as distinguished from racial evolution, are 
against this natural mandate regarding the family. The wisdom of British 
biologists, expressed by Tennyson in his memorable lines: 

So careful of the type . . . 

So careless of the single life, 

has been transmuted into the fatal reverse 

So careful of the single life 
So careless of the type. 

The closing decades of the nineteenth century and the opening decades 
of the twentieth have witnessed what may be called a rampant individual¬ 
ism—not only in art and literature, but in all our social institutions—an 
individualism which threatens the very existence of the family; this is the 
motto of individualism: let us obey our own impulses, let us create our 
own standards, let each individual enjoy his own rights and privileges—for 


4 


HENRY FAIRFIELD OSBORN 


tomorrow the race dies. In New England a century has witnessed the 
passage of a many-child family to a one-child family. The purest New 
England stock is not holding its own. The next stage is the no-child mar¬ 
riage and the extinction of the stock which laid the foundations of the 
republican institutions of this country. 

It is questions of this kind which are being set forth before this Congress 
so that they may be disseminated among our people. Let us endeavor to 
discard all prejudices and to courageously face the facts. Recent works by 
Bury and Inge on human progress are regarded in some quarters as pessi¬ 
mistic. I do not regard them as pessimistic, because to my mind the pessi¬ 
mist is one who will not face the facts, and these writers, especially Inge, 
look at the worst as well as at the best. I regard an optimist as one who 
faces the facts but is never discouraged by them. The optimist in science 
is one who delves afresh into nature to restore disordered and shattered 
society. This was the constructive spirit of Francis Galton, founder of the 
science of eugenics. I trust it will be the keynote of this Congress. To 
know the worst as well as the best in heredity; to preserve and to select the 
best—these are the most essential forces in the future evolution of human 
society. 
























• I: 

. 

■ 




























. 


































* 































Major Leonard Darwin 

Son of Charles Darwin, and president of the Eugenics Education Society of Great 
Britain, also head of the British delegation to the Second International Congress of 
Eugenics. 



THE AIMS AND METHODS OF EUGENICAL SOCIETIES 


LEONARD DARWIN 
President, Eugenics Education Society , England 

International congresses are organized no doubt mainly with the object 
of enabling workers in the same field both to become personally acquainted 
with each other—a far-reaching benefit—and to exchange information and 
ideas. We who have just crossed the Atlantic have come to a land in which 
many notable institutions have long been engaged in the study of biology and 
genetics, these being the pure sciences on which the applied science of eugenics 
is based, and where human racial problems have also long been keenly in¬ 
vestigated. So much has been done in all these directions here that when 
I was honored with an invitation to address you I felt great difficulty in 
selecting a subject which I could discuss with any reasonable prospect of 
promoting our common aim, namely, the improvement of the racial qualities 
of future generations. It is, however, not only scientific information 
which we can now profitably exchange one with another, but also our actual 
experiences; and, as I have been for ten years president of a British society 
for the promotion of eugenics, it occurred to me that it might interest you to 
hear something about our aims, our methods and our difficulties. I look 
forward to the time when eugenical societies will exist in all populous centers, 
their work being to strive to build up a social superstructure on the scientific 
foundations laid by central organizations engaged in biological and eugenical 
research. Whilst these much needed societies are passing through the 
period of their adolescence, we may be sure that they will not be without 
their growing pains and their difficulties; and these difficulties will certainly 
be more easily overcome if clearly realized in advance. I hope, therefore, 
that existing societies will not scruple to air their troubles in public! 

When an association is being created with any social object in view, a 
demand is likely to be made for a clear and rigid definition of the policy 
which is to be promoted by it; and from such demands may arise not only 
the first juvenile ailments of eugenical societies, but also occasional internal 
inflammations later in life. Now I was recently asked to state once again in 
broad and general terms what are the aims of my society, such a statement 
being needed not so much for our own information as to enable us to make 


5 


6 


LEONARD DARWIN 


our position more clear to the general public. The main difficulty in 
replying to this request lay in the fact that experience has taught us that 
attempts to decide in detail exactly what may be advocated and what should 
be condemned by eugenists are more likely to do harm than good by unduly 
restricting eugenic activities. A choice has always to be made between a 
smaller society with narrower aims and a larger society tolerating wider 
divergences of opinion; and although both plans have their advantages, yet 
in a young and growing subject like eugenics care should be taken not to 
injuriously hamper future liberty of action by too rigid definitions of 
policy. What seemed to me to be needed was a eugenic sign post, with 
arms pointing, not to every by-path, but to the various main roads along 
which our society should strive to advance; and the conclusions I then 
reached I now repeat in the hope that they may prove to be of some interest 
to a wider circle of friends. 

The first words which I uttered as the president of my society ten years 
ago were that heredity should be its guiding star, and in that opinion I have 
never faltered. A good deal of progress has been made since that date, and 
now the man who calls himself well educated is as a rule beginning to have 
^ome dim idea that all human beings are the product of two factors, heredity 
and environment, and that consequently to both of them some attention 
should be paid. Now if a eugenical society accepts only one of these 
factors, namely, heredity, as the foundation on which all its operations ought 
to be built, its members should as individuals most clearly emphasize the 
fact that all those who are striving to improve human surroundings have 
their warm sympathy. Of course eugenists cannot approve of such meas¬ 
ures as would injure mankind as a whole, the future as well as the 
present being taken into account; but, putting that possibility aside, we 
personally should give our blessing to many reforms which eugenical 
societies do not help to promote. We see as clearly as anyone that to take 
steps tending to produce in the future a race with the best possible natural 
qualities would be a futile proceeding unless we hoped that when such a 
race did appear great care would be taken to give to it good surroundings. 
If eugenical societies confine their attention exclusively to heredity, it is 
only because so many other societies think only of environment. 

It is true that sometimes it may be necessary to indicate that the high 
hopes entertained by reformers of to-day are not justified by past ex¬ 
periences. It may be said with only a microscopic divergence from the truth 
that all reforms since civilization began have been based on attempts to 
improve human surroundings; and we may ask those who found their 
hopes for the future only on changes being made in environment to con- 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


7 


sider how much has thus been accomplished since history began. As to 
our highest moral ideals, is it not true that for the most part they have been 
promulgated in certain eastern countries ever since the dawn of civiliza¬ 
tion? How do we compare in intellect with the inhabitants of ancient 
Greece two thousand years ago? With a knowledge of the delights of 
country life, can we look on our slums with anything but shame? Do we 
not blush to talk of peace on earth and goodwill towards men whilst remem¬ 
bering what has happened during the last seven years? And, in view of all 
this, have we any right to assume that improvement of environment will do 
more for mankind during the next two thousand years than it has done since 
the days of Plato? Reformers who look only to surroundings should con¬ 
sider well the foundations on which their projects are based before pointing 
the finger of scorn at the believers in heredity. Eugenics has been called a 
dismal science, but it should rather be described as an untried policy. 
Eugenics indicates a new method of striving for human welfare which, if 
combined with an equal striving for improvements in human surroundings, 
more truly justifies a hopeful outlook than anything which has yet been 
tried in the whole history of the world. More hopeful, that is, if the roads 
to which our eugenic finger post is pointing are not as studiously avoided in 
the future as they have been in the past. 

The eugenic sign post which we wish to erect should, in my opinion, have 
three arms on it, pointing to three main lines along which an advance should 
be pressed forward. In the first place the public should be made to realize 
more and more fully what a potent influence heredity has on the fate of 
all nations. In the second place efforts must be made to ascertain and to 
make known the rules by which each individual ought to strive to regulate his 
own conduct in regard to parenthood in accordance with the laws of heredity 
in so far as they are now surely known. Lastly, the action which the state 
should take in order to stimulate and to enforce conduct productive of 
racial progress must be considered, a line of advance to be advocated, how¬ 
ever, with great circumspection when compulsion is concerned. Our 
aim must be to advance along all these three roads simultaneously and 
continuously. 

The laws of natural inheritance supply a means of predicting in a measure 
the qualities of offspring when the qualities of their parents are known; and 
if any society accepts heredity, not as its sole guide, but as a light ever to 
be held in view, it is in fact intending to rely to some extent on these laws 
of natural inheritance when attempting to forecast the results in the future 
of our actions of to-day. Genetics is the pure science which deals with 
heredity, and genetics is, therefore, the very foundation on which the super- 


$ 


LEONARD DARWIN 


structure of eugenics is being built. The students of genetics will, how¬ 
ever, I am sure, all agree that a vast amount of research is needed before 
they will be able to rest satisfied with the knowledge they have acquired, 
supposing it to be possible that such a state of contentment will ever be 
reached. Now it is impossible to conduct the needed breeding experi¬ 
ments on human beings, and genetic research must be largely concerned 
with the lower animals and with plants; whilst eugenics is primarily con¬ 
cerned with man alone. Then again eugenics must include the study of 
many social and economic problems which lie quite outside the sphere of 
genetics. The pure science of genetics and the applied science of eugenics 
do, therefore, cover different fields, though the boundary between them is 
ill defined and movable; and in both fields further advances are urgently 
needed. For these reasons it seems to me—though here opinions may 
differ somewhat—that the main aim of eugenical societies should now be, 
whilst leaving geneticists to cultivate their own ground, to formulate a 
sound eugenic policy based on existing genetic knowledge, and then to 
promote the translation of every advance in eugenic theory into general 
practice. If we eugenists rely on scientific experts for the laying of our 
scientific foundations, then we shall be able to devote our main energies to 
the advocacy of reforms tending to promote racial progress and to con¬ 
sidering how wide may be the area over which such reforms can be justifiably 
extended. 

With regard to much of the research work which is so urgently needed, 
most eugenical societies will indeed have no option but to leave it to others 
or to leave it undone; because in many lines of enquiry a well equipped 
laboratory and a highly skilled staff are essential for success. Certain 
investigations, which need no special apparatus, however, could be carried 
on anywhere. Moreover, the scientific material as received from geneticists 
often needs to be thoroughly discussed by eugenists in a scientific spirit 
before being applied to human affairs; and we must not rely wholly on 
genetic research for the supply of scientific material on which to build. 
Wealthy patriots in all countries will doubtless from time to time perceive 
that by their wealth they might help to promote the acquirement of that 
knowledge on which racial progress must depend in the future. A strong 
central society might in such cases play a useful part in suggesting various 
directions in which, with their aid, advances of great value could at once be 
made; as well as being ready, if so desired, to act as agents by whom the 
investigator would be selected and employed, care being taken not to ham¬ 
per him with undue control. The more liberal the benefaction the more 
fundamental and far-reaching might be the researches thus undertaken, 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


9 


and the greater the ultimate benefit to mankind. Your endowments in 
America are so magnificent that you may not fully perceive how much they 
are needed elsewhere. 

As to the first of the suggested lines of advance, namely, as to getting into 
direct and immediate touch with the public with the hope of spreading 
abroad a general knowledge of the laws of natural inheritance, this knowl¬ 
edge should form the basis of all the arguments brought forward at public 
lectures on eugenics, that is, at lectures not forming part of any ex¬ 
tensive series. It is indeed in laying this foundation of scientific truth 
that speakers on such occasions encounter their greatest difficulties; for 
many prejudices arising from ignorance have to be overcome. For ex¬ 
ample, those who do not acknowledge to themselves that men differ 
greatly from each other in their inborn qualities, cannot be made to realize 
the extreme importance of paying attention to heredity in regard to 
social questions; and the acknowledgment that we do not start even in 
the race of life will be hindered by a disinclination which we all feel 
both to regard any human disabilities as being incurable and to own that 
other individuals may be greatly superior to ourselves. As to the facts 
on which the scientific theories of heredity are based, it is worse than useless 
to attempt to give them in detail at single lectures; for lecturers should 
remember that on such occasions they cannot hope to do more than leave an 
enduring general impression on the minds of their audiences. Except in 
systematic courses of study, much must always be both stated and accepted 
on authority; for to fully justify all the beliefs of eugenists would require 
months rather than days. “It is hardly possible,” so my father declared, 
“within a moderate compass to impress on the minds of those who have not 
attended to the subject, the full conviction of the force of inheritance which 
is slowly acquired by rearing animals, by studying the many treatises which 
have been published on the various domestic animals, and by conversing 
with breeders .” 1 If this be so, the public can only learn how to give to 
natural inheritance its proper value by acquiring information at second 
hand; and yet to make any statement acceptable to audiences, it must be 
in some degree endorsed by their own reasoning powers. It is on this 
account that allusion to the breeding of domestic animals becomes almost a 
necessity in public lectures on eugenics, for the wisdom of attending to breed 
in the case of cattle and dogs is universally admitted. Great care should, 
however, always be taken to indicate that, though our experiences in the 
stockyard enable us better to understand the laws of natural inheritance, 

1 Animals and Plants under Domestication, Darwin I, pp. 447-448. 


10 


LEONARD DARWIN 


yet our reliance on these laws carries with it no implication whatever that 
the methods of the animal breeder ought to be introduced into human 
society. It should in fact be most strongly emphasized that nothing which 
we advocate is contrary to the highest religious ideals. This is, however, 
rather a digression; for I am not here to instruct lecturers how to lecture. 
All that I now wish to insist on is that, by means of lectures to audiences of 
all kinds, the endeavor to spread abroad sound impressions concerning the 
force of natural heredity and the enormously important influence which it 
has in deciding the welfare and the destiny of nations should form a promi¬ 
nent part of the programme of all eugenical societies. 

The title selected for the British Society by its founders was the Eugenics 
Education Society, and certainly they had excellent reasons for thus empha¬ 
sizing the educational aspects of the eugenic campaign which they were 
inaugurating in my country. No class of the community is more im¬ 
portant to interest in racial problems than teachers of all grades; because 
the ideas of the youth of to-morrow will depend so largely on the opinions 
of the teachers of to-day. But teachers must be taught before they can 
take a thoroughly intelligent interest in racial questions; and for this 
reason it is of primary importance that biology should be given adequate 
recognition in the curricula of all colleges where teachers are trained. Our 
educational aspirations could not, however, be completely satisfied in this 
way; for to finally succeed in the first of our main aims, namely, the spread¬ 
ing abroad of a general knowledge of the laws of natural inheritance, 
natural science must be given a far more prominent place than at present 
in the courses of studies of all schools and colleges. No doubt there are 
many who now regard our efforts with great distrust; but those who feel 
thus should remember that the better and the more wide-spread the teach¬ 
ing of biology, the more certain would it be that any eugenic errors would be 
detected and their harmful influence prevented. Moreover, if we want 
progress in scientific research to be both rapid and on right lines, it is 
important that a considerable number of students should be thoroughly 
trained each year in genetics, or that more undergraduates should specialize 
in natural science at our universities than at present. Eugenics has a long 
struggle before it, and all these methods of laying educational foundations 
for future progress should certainly come within the scope of the efforts of 
eugenical societies. 

Passing on to the second of the main lines along which eugenical societies 
should strive to advance, what we want to know is the rules which ought to 
guide each individual in deciding on his own voluntary actions in all matters 
relating to racial progress. The attempt to ascertain the precepts by means 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


11 


of which each one of us should strive to regulate his conduct in questions 
connected with parenthood obviously involves the consideration of a number 
of ethical, racial and economic factors; for, in regard to any proposed line of 
conduct, we have to weigh in the balance as well as we can its moral effects, 
the immediate material advantages or disadvantages to the family and to 
the state which are likely thus to arise, and the benefits or injuries which 
it will confer or inflict on the race in the future. Even if these problems be 
approached in a calm and scientific spirit—and in this respect eugenical 
societies should strive to set a much needed example—even then it will be 
exceedingly difficult in most cases now to arrive at precise conclusions. 
We must not attempt in the present state of our knowledge to lay down 
rigid rules of conduct, but only to suggest general guiding principles; 
though we may hope that with every advance of science it will be possible 
more and more clearly to indicate what each individual ought to do and 
what he ought to avoid. As an illustration of the difficulties involved in 
these problems, consider the case of a contemplated marriage when both 
families thus to be connected are characterized by some degree of ill health. 
Now it would only be persons endowed with high moral qualities who 
would be likely to obey any self-denying ordinance in regard to marriage 
and whose fertility would, therefore, thus be diminished. Might we not, 
by condemning marriage in such cases, tend to breed out the most valuable 
of all human attributes, namely, the desire to do right? Again if insanity 
were the family trouble in question, this being one of the most grievous of 
all human ailments, we now know that it is sometimes the result of disease 
and probably in such cases not heritable, whilst other types certainly do 
run in families. What are we to do in the face of such doubts and difficul¬ 
ties as these? Are we to admit our incapacity to meet the situation? 
Certainly not, for the history of scientific research clearly proves that what 
to-day appears like an impenetrable barrier to further progress will probably 
to-morrow be regarded rather as a useful stepping stone for a further ad¬ 
vance. Doubtless we have difficulties ahead of us, which must be faced 
with patience; but we should take note of these obstacles in our path mainly 
as emphasizing the need for societies where such guiding rules for voluntary 
conduct in relation to parenthood as are warranted by existing knowledge 
and by present needs will be wisely and temperately discussed. 

A comparatively new subject like eugenics is apt to arouse prejudices and 
to give opportunities for misapprehension; and it sometimes seems that 
what is now most needed on the part of eugenical societies in regard to 
voluntary actions is that they should make clear what they are not recom¬ 
mending. We have been accused of wishing to abolish love altogether as a 


12 


LEONARD DARWIN 


guide to conduct; but this is false. What we desire is rather to purify love, 
or to clear away all those harmful influences which so often attach them¬ 
selves to it. Certain American investigations indicate that the ideals 
which naturally dwell in the minds of young people in regard to the qualities 
of the mates to whom they would wish to be connected in marriage are on 
the whole fairly sound, and that these promptings if followed would gener¬ 
ally lead to unions beneficial to the race. But the desire for wealth, the 
wish to rise in the social scale, and, some would add, too great attention to 
personal appearances, often make the choice of a mate far worse than it 
would have been if these natural ideals had been given full sway. In 
passing I must, however, put in a racial plea for good looks on the ground 
that they are apt to be associated with good health; a plea which I hope 
does not spring from a mere masculine weakness on my part. Be that as 
it may, love is doubtless to a large extent aroused by advantageous moral 
and mental qualities; and, in so far as that is the case, it forms the firmest 
foundation on which to base a eugenic policy. Much can be done to 
help to lay this foundation by promoting suitable opportunities for the 
meeting of young men and maidens; by judiciously encouraging intercourse 
between our children and worthy friends of the other sex, from amongst 
whom worthy mates are not unlikely to be selected; by stimulating>a pride 
of family in so far as dependent?on character and performance; and, above 
all, by fostering the growth of all that is noble in the ideals of the adolescent. 
Never make a close friend of a person one can not respect is, I believe, 
not only a helpful rule of life, but also a useful way of setting an example 
to the rising generation. But here a possible racial danger must be noted; 
for an injudicious pursuit of the policy here suggested might make the high- 
minded become too particular and therefore less likely to marry than their 
more ordinary companions, with obvious dysgenic consequences. Pure 
love between the sexes should be proclaimed as the noblest thing on earth, 
and the bearing and rearing of children as amongst the highest of all human 
duties. Some risks ought to be run in order to secure these joys and to 
fulfil these duties; and Cupid may well remain a little blind to all minor 
defects. To promote these ways of regarding sexual problems and to show 
how often the moralist unknown to himself is in effect striving to better the 
racial qualities of future generations come well within the scope of our 
endeavors. 

Though we have seen that as knowledge increases so the difficulties of 
deciding on rules of personal conduct will diminish, yet it is certain that 
these difficulties will ever remain very formidable. We may now boldly 
assert that when the heritable defects of many members of a family are 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


13 


very serious, those belonging to it should not become parents; but how 
serious must these defects be before being regarded as a bar to parenthood? 
It will never be possible to draw as sharp a line of demarcation as that 
between sheep and goats when marking oil from the general population 
those in whom parenthood would be a moral offense. Because of this 
impossibility, it may come to be held that the size of the family should 
vary with the innate qualities of the parents; but how is this relationship 
between fertility and transmissible characteristics to be determined? 
Then, again, many who take no thought concerning racial questions now 
hold strongly that it is wrong to bring a child into the world without a 
reasonable prospect of its being able to live a life up to a certain standard of 
civilization. But what should be the standard adopted? In large numbers 
of cases the cause which has prevented the winning of a “standard” liveli¬ 
hood, however we may define that term, has been some inborn defect, or 
defect which would in a measure be passed on to the next generation. 
Teach those not living up to standard to regulate their conduct with due 
regard to the welfare of any children who may or may not be born in the 
future, and many would limit their families on this account; with the 
results that these harmful innate defects would appear less frequently in 
future generations. Is it not, therefore, of great importance that some 
attempt should be made to ascertain what standard of living does justify 
parenthood? Again it is even more important that it should be widely 
felt that it is morally wrong to limit unduly the size of the family when 
parents are up to “standard” in all respects; for it is essential for the 
welfare of mankind that the seed of this good stock should not be lost to 
posterity. Eugenical societies should, in my opinion, steadily keep in view 
the necessity of trying to solve all these intensely difficult problems; prob¬ 
lems which need the joint consideration of the eugenist, the geneticist, and 
the economist for their solution. But as for our advice of to-day con¬ 
cerning personal conduct in regard to procreation, we can say little more 
than that moral principles must always be kept in the foreground, and 
that, for the rest, trust must be placed in common sense and a wise doctor . 2 

To whatever extent success may attend our efforts to lay down rules for 
personal conduct in regard to parenthood, to that extent we shall have 
succeeded in deciding on the directions in which we wish to advance in 
these matters. Such decisions will, however, prove to be but a very un¬ 
certain indication of the extent to which the state should endeavor to 

2 1 assume that the doctor has studied genetics, which is unfortunately not always 
the case. 


14 


LEONARD DARWIN 


promote or to enforce obedience to these rules; this being the subject to 
which we must now turn our attention. By promoting uniformity of 
conditions and by checking individual initiative, the state often retards 
progress; and, besides affecting those intended to be affected, govern¬ 
mental action nearly always produces on other persons various consequences 
which were unforeseen and which are never fully realized. Whatever may 
be our political opinions, we nearly all of us agree that these are dangers 
which must be taken into account when contemplating state control over 
the individual. These are, however, large issues which some will regard as 
lying outside the proper scope of eugenic considerations; whilst the point 
which I especially wish to emphasize in this connection is one definitely 
related to the actions of eugenical societies. In my opinion our societies 
ought to be ready to encourage discussion on all proposals for relevant 
reforms, whilst they should be cautious in the present state of our knowledge 
in actually recommending governmental interference. If discussion be not 
bold, progress will be slow; for a nation can not grope its way quickly to 
the front in the darkness of ignorance. If action be too bold, progress will 
also be slow; for the wrong road will often be taken. In matters of conduct 
we should balance the probability of good or evil arising from the action 
proposed to be taken, as against the magnitude of the good or evil if it does 
arise. The smaller the chances of failure, the smaller may be the benefits 
hoped to be attained. The probability of harm resulting from the mere 
discussion of any reform would usually be very small, even if that reform 
would be very harmful if adopted. On the other hand, the possibility of 
benefits arising from the discussion of reform is almost equally obvious 
whether the proposed legislation would in fact be beneficial or harmful. 
To take a single example, there are strong differences of opinion as regards 
sterilization; but all may hold that by open discussion true conclusions 
would most likely be reached. The advocates of sterilization of course wish 
to have this subject brought to the notice of the public; whilst its opponents 
must admit that they will be more likely to promote than to retard its 
introduction, by, as it were, burying their heads in the sand like the ostrich 
and by refusing to favor the creation of opportunities for openly stating 
their objections to it. It is indeed nearly true to say that every subject 
may be openly discussed with advantage provided the occasion be properly 
chosen and it is in this spirit that eugenical societies should, in my opinion, 
conduct their proceedings. 

In all human affairs we are constantly being compelled to take opposing 
considerations into account and to adopt compromises, and I think that I 
ought not to be accused of inconsistency if I now turn round and show why 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


15 


eugenical societies ought not to be too timid in regard to legislation. As 
to your middle-aged Anglo-Saxon, and I am only speaking for my own 
country, there is hardly anything which he dislikes so much as having to 
change his opinions; and from this weakness men of science are by no means 
exempt! Here is a barrier which will stop any half-hearted advance on 
the part of eugenic reformers! To the students of natural sciences, at all 
events, we can suggest that Nature’s plan seems to have been to stamp 
out of existence all organisms which fail to fill the places she assigns to 
them, and this without regard to the sufferings thus caused or to the 
superiority in many respects of large numbers of the individuals thus 
eliminated. By adopting rational methods in human affairs, much can be 
done and much ought to be done to prevent human beings from being 
enforced to sufferings similar to those which animals in the wild have to 
endure because of that struggle for existence to which they must submit; 
but nevertheless we should not be quite blind to the example set us by 
Nature in her readiness to sacrifice the individual for the sake of the race. 
Unfortunately it will be our politicians who will mainly settle how far the 
teachings of science shall be made to affect legislation; and this they will 
be apt to do with little reference to the opinions of experts and largely in 
the hope of catching votes. But the votes of future generations can not 
now be caught, and their interests will, therefore, be likely to receive but 
scant attention in all democratic countries. Governments which depend 
on the suffrages of the people are of necessity always somewhat timid in 
regard to unpopular reforms; and until eugenics becomes popular—when 
will that be, I wonder!—there is not the slightest chance of eugenic reform 
moving forward with too rapid strides. Eugenists must lead the advance 
in racial questions, and our societies must remember that nothing is more 
fatal to leadership than a show of timidity. We should discuss long and 
freely, and when we do advance, advance boldly. 

Legislative reforms can seldom be effectively promoted or steadfastly 
maintained unless they are sanctioned by the general opinion of the citizens 
concerned; and, on somewhat similar grounds, eugenical societies would be 
wise to avoid taking corporate action in regard to legislation unless the 
proposal in question has the nearly unanimous approval of their members. 
The neglect of such warnings has led to the disappearance of governments 
and to the disruption of societies! When legislation does not involve com¬ 
pulsory interference with the liberty of the individual, there is comparatively 
little danger of internal friction being caused by its advocacy; for unanimity 
in such circumstances is both more probable and less necessary than when 
compulsion is involved. As examples of legislation of general application 


16 


LEONARD DARWIN 


producing beneficial racial effects, certain reforms in regard to taxation 
might be mentioned. My Society took an active part in the agitation in 
favor of such alterations in the assessment of income tax as would make the 
burden of taxation fall less heavily on parents of families and more heavily 
on bachelors and the childless in the same stratum of society , the object being 
to increase the birth rate of a useful class of the community. As to legisla¬ 
tion involving interference with individual liberty, here also unanimous 
support can be obtained if the racial advantages are sufficiently obvious. 
For example, there was no dissension whatever in my society when we moved 
in favor of the Mental Deficiency Bill, a bill which authorized the segrega¬ 
tion of the feeble in mind, that is to say, their detention in comfort under 
carefully safeguarded conditions. But until unanimity in the ranks of a 
eugenical society in regard to such compulsory measures is obtainable, their 
discussion only is to be recommended. Personally I should like to see 
practical steps at once taken for lessening the fertility of habitual criminals, 
of hopeless wastrels, and of the grossly unfit generally, and others doubtless 
wish to advance in other directions; but we must have patience. My 
object for the moment is not, however, to attempt to survey all the roads 
by which advances may be made in future, but rather to consider what 
should be the broad principles of strategy which should guide eugenical 
societies in the long fight before them in their attempts to promote racial 
progress. 

Thus I have dealt with the objects which eugenical societies should strive 
to attain rather than with the methods of attaining the ends desired, the 
reason being that I have little novel to suggest in regard to methods. 
With the view to the advancement of scientific knowledge and the elucida¬ 
tion of eugenic problems, my society holds periodical meetings at which 
addresses are delivered or questions debated. In our Review these ad¬ 
dresses are often published, and we there also try to give impartial ac¬ 
counts of current eugenic literature. We maintain a library, and give 
advice to readers. We keep in touch with foreign societies, and it has been 
an especial pleasure to us to give all the assistance in our power to the 
American committee which has so admirably organized this Congress. 
As to activities definitely undertaken for the purposes of propaganda, the 
following may be mentioned: the delivery of lectures to audiences of 
various types, including social clubs, debating societies, educational con¬ 
ferences, summer schools for teachers, and, during war times, soldiers in 
camp and barracks; the organization of summer schools dealing largely with 
eugenics; the sending of deputations to government departments; and of 
letters to the press. To take one example in detail, after a thorough enquiry 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


17 


concerning the incidence of our income tax, a letter was written to all 
members of Parliament, and at a later stage amendments to the Finance 
Act were proposed by members at our suggestion, and were rejected I 
The next step, a direct result of this agitation, was the appointment by the 
government of a royal commission on the income tax before which I gave 
evidence on behalf of my society. Several of the recommendations of 
that commission, representing a step forward in the direction desired, were 
subsequently adopted and became law. Thus by steady persistence on well 
thought outlines a society may be able to produce material effects in many 
directions. As a last word about the doings of my own society, I must be 
allowed to mention a dinner followed by an address, held on February 16 
in each year. In this way we yearly remind ourselves on the birthday of 
Sir Francis Gal ton that to him we owe the opening of the eugenics campaign 
in England. 

What I have tried to do in my address today has been to give some indica¬ 
tion of the difficulties likely to be encountered by youthful eugenical 
societies; difficulties which, we have seen, may come from many quarters 
and in many shapes. Questions connected with both sex and personal 
liberty have to be dealt with by eugenists, and these are topics especially 
liable to give rise to strong feelings. Even when the opposition thus 
aroused is quite unreasonable, we should, however, always remember that 
the sentiments underlying this opposition are often in many respects highly 
commendable, and that to openly acknowledge where others are in the right 
is often the best way of getting a hearing for ourselves. The most formid¬ 
able foe we have to meet is ignorance; and here again it is wise to admit that 
the ignorance is not all on one side. With every growth in our knowledge of 
biology and sociology we shall be able safely to enlarge our programme, and 
we should make it clear that our discussions of to-day are often tentative 
and do not always indicate the directions in which we shall advance to¬ 
morrow. As to the ignorance of our opponents, it can only be overcome by 
patience, perseverance and above all by never concealing such doubts as 
are still felt. Unfortunately it must be admitted that even perfect knowl¬ 
edge, however widely held, would not make our path quite smooth, human 
nature being what it is; for the want of attractiveness of our programme is 
largely due to the fact that we are looking to human welfare in the more or 
less distant future and not to present-day comforts. Most men in their 
march through life are hoping either for personal distinction as a reward 
for their exertions or for quick returns on their investments; neither of 
these benefits is to be obtained in the eugenic market. You can easily 
enough get your forests cut down and the timber sold for an immediate 


18 


LEONARD DARWIN 


profit; but the planting of slow growing trees, which will not be worth 
felling till most of us are dead, is a less attractive venture, though more 
beneficial to the nation. The reforms which the eugenist wishes to plant 
would certainly bear excellent fruit in due course, even though much of it 
would only be gathered by our children and our children’s children. Then 
again your business men not seldom try to sell their goods by running down 
the wares produced by their rivals, an inexcusable proceeding in so far as 
merely an outcome of greed and jealousy. Now this same competitive 
spirit is far too much felt in social work, and I fear we eugenists have often 
aroused opposition by unnecessarily running down reforms dependent on 
changes in environment. Let us rather strive to show that there is plenty 
of open ground over which reformers of all kinds can strive to advance 
simultaneously and harmoniously; and let us all recognize that jealousy 
is one of the commonest and probably the most insidious of all human 
failings. The claims of this generation and of posterity are doubtless 
sometimes antagonistic, and the genuine difficulties thus arising must be 
openly faced and often met in a spirit of wise compromise. The main 
obstacles to be overcome by eugenists are, however, dependent on moral 
failings, and what we have to show is that we are engaged in a moral cam¬ 
paign, with human welfare in the highest sense as the goal for which we are 
striving. 

Eugenics aims at increasing the rate of multiplication of stocks above the 
average in heritable qualities, and at decreasing that rate in the case of 
stocks below the average. But if the banner under which we are to fight 
should only have inscribed on it some such arid definition of policy as this, 
our defeat would be certain. We must prove that we are under the 
guidance of a noble ideal. We of this generation are responsible for the 
production of the next generation and, therefore, of all mankind in the 
future; and all in whom this sense of racial responsibility acts as a deep- 
seated sentiment, greatly affecting their action and their policy, are in 
truth guided by the eugenic ideal. The belief that man has been slowly 
developed from some ape-like progenitor came towards the close of the last 
century to be nearly universally held by thoughtful persons; this belief 
gave rise to a new hope that this upward march of mankind might be con¬ 
tinued in the future; and out of this new hope sprang the eugenic ideal. 
This growing understanding of the past history of the world has led us to 
see that, if we are to imitate Nature in her methods, we must be content to 
advance by means of a long succession of small steps; just as rain falling in 
drops on the earth has slowly carved out mighty valleys in the hardest 
rocks. Without constructing wild Utopias, we must be content if some 


AIMS AND METHODS OF EUGENICAL SOCIETIES 


19 


little racial progress can be ensured as each generation succeeds another; 
for to work in this spirit is to work in harmony with the knowledge which 
gave birth to the eugenic ideal. Progress on eugenic lines will make man¬ 
kind become continually nobler, happier, and healthier; whilst those who 
imagine that our sole aim is to make man a stronger animal or a better 
beast of burden are utterly ignorant of the meaning of the eugenic ideal. 
But science, whilst giving us good grounds for hope, also issues a grave 
warning concerning the danger of national deterioration resulting from 
the unchecked multiplication of inferior types. In the past many nations 
of the first rank, when apparently advancing without check on the path of 
prosperity, have begun to decay from unseen causes, and have in time so 
fallen from their high estate as to cease to count as factors making for 
progress. A determination that such a downfall shall not be the fate of his 
nation is a sentiment felt by every man who is animated by the eugenic 
ideal, an ideal to be followed like a flag in battle without thought of 
personal gain. 


RESEARCH IN EUGENICS 


CHARLES BENEDICT DAVENPORT 
Carnegie Institution of Washington 

Man is studying all phenomena. He has at last come to study himself. 
Not his diseases, not his language, not his customs merely, but also his more 
intimate self. Man is studying man as an animal, who varies in his traits, 
who selects his mates for better or worse, who has a larger or smaller 
number of children that are more or less healthy and live for a varying 
period. The races of man are being studied not merely to list their 
differences, but to find how those differences arose and how they are 
transmitted to progeny and how they intermingle. We are studying the 
laws that govern the distribution of traits in the family; we are studying 
the consequences of combinations of these straits in the instincts, interests 
and behavior of individuals. At last we are studying man as the product 
of breeding and as the subject of an evolutionary process. And we are 
studying the human germ plasm, its composition, its mutations and its 
mixtures. * 

And why do we investigate? Is not enough known to warrant propa¬ 
ganda; and should we not better organize for a campaign to change what 
needs changing? Alas! we have now too little precise knowledge in any 
field of eugenics. We can command respect for our eugenic conclusions 
only as our findings are based on rigid proof, a proof that is either statistical 
or experimental. Only as we are able to base our statements on scientific, 
quantitative data can we hope to carry conviction and not arouse con¬ 
trary opinion. People do not have heated discussions on the multiplication 
table; they will not dispute quantitative findings in any science. 

It is largely due to the extraordinary vision of Mrs. E. H. Harriman, the 
founder of the Eugenics Record Office, that in this country eugenics is more 
a subject of research than of propaganda. She maintained that we should 
be more concerned with knowing than with doing. Ascertained facts do 
not require propaganda. 

It is sometimes asserted that research in eugenics belongs to the realm of 
applied science, and much of it does. But not all. There are fields of 
eugenical research, especially in human genetics, that are pure research in 


20 


RESEARCH IN EUGENICS 


21 


as much as they are devoted to investigations that can not be carried out 
so well on any other material. For example, inheritance of psychological 
traits, of temperament and of sense perception. 

In so far as eugenics may lay claim to being a science, it has not only a 
subject matter—but also a method of its own. In studying the genetics of 
the lower animals, we proceed by the method of control of matings. Now 
this method is obviously not applicable to man in modern civilized countries. 
It has to be replaced by the collection of the history of matings that have 
been already made and a study of their progeny. We replace the experi¬ 
mental mating of the geneticist with the principle that every fertile human 
mating is an experiment in genetics, and it is for us to record the result of 
the experiment. Some day, we may hope, human matings will be carried 
beyond the stage of experiment. 

At present, then, the student of human genetics must collect data on 
human matings and their outcome. Of course, he must know, as thoroughly 
as he can, the genetic nature of the matings; so that he can give the probable 
genetic composition of the gametes. This means that he must know for the 
mated pair, the parent, uncles and aunts and their children. He can then 
check his findings by studying the traits of the children. Since the capacity 
of one man for collecting by himself is very limited, it is necessary to train 
observers to collect data. Hence has arisen the profession of eugenical 
field worker whose function it is to study through three or more generations 
and as analytically as possible all the members of an inter-generating group 
so that their probable genetic composition may be known. By gathering 
together in one depository a large quantity of carefully ascertained family 
dala, the basis is laid for human genetical studies. 

/The history of the development of the method of eugenical field workers 
is not a long one. Dr. Alexander Graham Bell was one of the first to use it 
extensively in this country. He employed such field workers in his study 
of deaf mutes especially those of Martha’s Vineyard, in the early eighties?/ 
The Reverend Oscar McCulloch made use of field workers in his study of the 
Ishmaelites in the nineties, and at the Vineland Training School such 
workers were employed before 1910. A large number of eugenical field 
workers (about 200) have been trained by the Eugenics Record Office since 
its beginning in 1910. 

Besides trained field workers, numerous volunteers are in a position to 
contribute data. Thus, in 1884, Francis Gal ton distributed his question¬ 
naires called “Record of Family Faculties,’’and over 150 persons volunteered 
to fill them out and return them to him for study. The Eugenics Record 
Office has made use of a similar questionnaire called “Record of Family 


22 


CHARLES BENEDICT DAVENPORT 


Traits,” of which 4,000, of varying degrees of excellence, have been deposited 
in that office. Some of these “Records” are exceptionally valuable. It 
appears that many persons show the capacity for and interest in filling out 
such schedules excellently. A few others will take the pains to make a still 
more detailed analysis of the individuals of their families. Many of these 
records have to be considered as finders merely; as guides to further 
inquiries. 

Additional records that are often of value are the printed genealogies 
and town histories, of which so many have been printed in this country, 
especially for the northeastern section. In addition, biographies, especially 
sets of biographies relating to members of a single family, will yield to the 
analyst of human traits data of the greatest importance. Finally all 
records—those of field workers, of volunteers and the printed records—must 
be indexed by name, place and trait so that their contents shall be readily 
available. 

In inquiries into human genetics it is desirable, where possible, to breed 
experimentally mammals, if any are available, which show the same trait 
that we are studying in humans. This is often possible, and such study will 
afford a control of results gained on man. Thus have been studied hare-lip 
in dogs, fecundity in sheep, instincts in dogs, polydactylism in fowls. 

In other studies the method employed will be that of accumulation of 
statistics, their tabulation and analysis. Thus we investigate mate selec¬ 
tion, the relative fecundity and relative mortality of the various stocks and 
the effect on the germ plasm of a country of the different immigrant races. 

Some of the results of analytical study of these eugenical data are fairly 
well established. A few clearly simple Mendelian traits have been found. 
Such is eye color in which brown is dominant over its absence. It is 
possible that in some cases additional factors may be present, but the rule 
serves as a first approximation. Dominant, also, appears to be curliness 
of the hair as contrasted with recessive straight. And there are various 
diseases and defects that appear either as simple dominants or recessives, 
such as abnormalities in number and form of fingers and toes, which are 
mostly dominant over the normal condition; various defects of the eye such 
as cataract, certain types of congenital deafness, various abnormalities of 
skin, and hair and nails. 

Other, and probably many other, traits are due to multiple factors—so 
often this is true as to suggest the hypothesis that in mammals, as contrasted 
with insects, traits are genetically relatively complex. Thus stature and 
build and proportions of parts and pigmentation of hair and skin are 
dependent on multiple factors. Indeed, there seems to be evidence that 


RESEARCH IN EUGENICS 


23 


negro skin color is dependent upon two pairs of factors which merely 
reinforce each other. 

Other traits are associated with sex in the remarkable fashion called 
sex-linked. That is, they are usually found only in the male sex and are 
inherited through the mother, though she, herself, is not affected. In such 
cases one usually finds male relatives of the mother who are affected. 
Such are color blindness, hemophilia and atrophy of the optic nerve. The 
facts of sex-linked heredity bring home, even to the layman, the lesson that 
heredity is a matter of the gametes; and that bodily appearance often gives 
no hint of the nature of the particular germ cells carried and, in so far, of 
what the inheritance shall be. The parents of an albino may have pig¬ 
mented hair and skin, but both carry gametes which lack the capacity of 
forming pigment. 

Our knowledge of the inheritance of these physical traits is sufficiently 
precise to be applied practically in cases of doubtful parentage. If the 
child, the known mother and both of the putative fathers can be seen, and 
some inquiry be made as to family stock of the three adults a decision can 
generally be rendered with a high degree of certainty ranging from 75 to 99 
per cent. For usually there will not be one critical trait merely but several 
traits whose combined evidence will be overwhelming. Already the 
Eugenics Record Office has been asked to answer certain questions about 
the inheritance of traits in a case of a claimant who maintained that he was 
the son of a wealthy man who died without known heirs. As lawyers get 
more used to the idea of utilizing the advances of knowledge for evidence, 
it is probable that eugenical knowledge will be more and more called upon. 

Not only of the physical traits referred to above but also of those of 
behavior we are learning the hereditary basis. It appears probable, from 
extensive pedigrees that have been analyzed, that feeble-mindness of the 
middle and higher grades is inherited as a simple recessive, or approximately 
so. It follows that two parents who are feeble-minded shall have only 
feeble-minded children and this is what is empirically found. It has been 
urged against this finding that it is improbable that so complicated a thing as 
full mentality depends upon only one factor. On the other hand, a con¬ 
sideration of the effect of internal secretions, of thyroid, of hypophysis and 
others leads to the conclusion that a brain with well differentiated in¬ 
tellectual centers may fail of complete development because of the absence 
of proper developmental impulses of glandular origin. Two persons whose 
brains are thus under-developed may differ greatly in their mental capaci¬ 
ties, because they have fundamental nervous differences, just as seedlings 
of different species, while all alike under-developed, differ in certain specific 


24 


CHARLES BENEDICT DAVENPORT 


traits. Apparently one group of hereditary mental defectives is such 
because those who belong to it lack a single factor for an adequate develop¬ 
mental impulse. 

Epilepsy, of the ordinary juvenile, dementing type, seems to be due, 
like feeble-mindedness, to a single developmental defect. Also, dementia 
prsecox has been found by several investigators to be due to a similar cause. 

But not only mental but also emotional states have a hereditary basis. 
The prevailing depressed mood appears to be due to a glandular condition 
that is determined by a certain developmental defect; and a prevailing 
excitability appears to be determined by a hereditary condition, which may 
be a tendency to excessive secretion of the suprarenal glands. 

Moreover, the quality of our senses has a clear hereditary basis, as the 
still unpublished work of Dr. Hazel Stanton on musical families clearly 
shows. It appears from these studies that not only have great musicians 
an innate capacity for discriminating between closely similar qualities of 
pitch, intensity, time and for tonal memory but they belong to families 
with these innate capacities. Also, it has been shown that these capacities 
are not improvable by training; they depend upon our very constitution. 
Now we have evidence that persons who have these capacities enjoy exer¬ 
cising them. Those in whom the capacities are slightly developed get no 
pleasure from exercising them. We conclude that the reason why musical 
people are such is primarily because of their possession of inborn musical 
capacities. The musician is born, not made. From these principles 
certain deductions seem naturally to flow. A great color artist is one in 
whom the innate capacity for color discrimination is well developed and 
his family shows other examples of colorists. The sculptor has the heredi¬ 
tary capacity for form discrimination and that is why he finds his highest 
pleasure in the art. The author is one whose verbal machinery is especially 
perfect. The sailor is one who finds his greatest pleasure in the beauty of 
form of the vessel, or perhaps in broad horizons and distant lands; he is 
neither claustrophil, nor domestic. In general, our vocations, or at least 
our avocations, are determined by our sensory structure and this is 
hereditary. 

The fact that not only our physical but also our mental and tempera¬ 
mental characteristics have a hereditary basis has certain important social 
bearings. It leads us to regard more charitably the limitations of our 
fellow men. The false doctrines of human equality at birth and of freedom 
of the will have determined a line of practice in the fields of education and 
criminology that, it seems to me, is not productive of the best results. 
In education we must know the child’s native capacities before we can 


RESEARCH IN EUGENICS 


25 


properly train. In dealing with delinquents we must know the hereditary, 
mental and emotional make-up before we can get an explanation of the bad 
conduct and before we can intelligently treat the delinquent. Organized 
society is too prone to “pass the buck” of its own shortcomings to the 
hypothetical “bad-will” of the offender against the mores. We should do 
better if we treated the misdemeanant as we treat a puppy whose actions 
displease us. Either train him carefully, if he is trainable; otherwise, put 
him in a position where the exercise of his instincts will not offend us. 

The relation of the glands of internal secretion, commonly known as 
endocrine glands, to human development and human behavior is becoming 
daily more obvious. Stature, build, proportions; details of development of 
bone, teeth, nails, hair, skin; intelligence, emotional control, all these 
things can be shown to be influenced by endocrine secretions. Indeed, it 
seems naturally to follow that the hereditary differences between people are 
due to hereditary differences in the activity of these glands. Now these 
glands, as is well known, secrete substances called “hormones” which 
regulate our physical, mental and temperamental constitution. The 
special quality and quantity of these hormones is determined by the idio¬ 
syncrasies of the enzymes of the germ cells. The hormones that determine 
our personality, constitute the bridge that connects this personality on the 
one hand, with the specific enzymes packed away in the chromosomes of the 
germ cells, on the other. You and I differ by virtue of the difference of 
atomic structure and atomic activity of the enzymes and hormones which 
make up that part of the stream of life-yeast which has got into and is 
activating our protoplasm and will activate that of the fertilized egg that 
results from us and our consorts. Thus each is what he is in his physique, 
in his thoughts and in his reactions largely by virtue of the peculiar prop¬ 
erties of those extraordinary activating substances, which are specific 
for him and other members of his family and race or biotype. The future 
of human genetics lies largely in a study of these activities, and the origin 
of differences or mutations in them. 

The study of human genetics leads into numerous fields of the physiology 
of human reproduction. Of these one of the most significant is that of 
twin-production. This topic has many aspects. As is well known twins 
are of two types. Two-egg twins come from two eggs simultaneously 
ovulated and one-egg twins arise by a division into two embryos of a single 
young embryo. The two children which thus arise from one egg are often 
so marvellously similar that they are called “identicaltwins.” Now these 
identical twins give a measure of the relative importance of heredity and 
environment, as Francis Gal ton pointed out. It is, indeed, marvellous to 


26 


CHARLES BENEDICT DAVENPORT 


see how such twins, even though living far apart, retain their initial resem¬ 
blance, experience at almost exactly the same time similar disease and 
emotional disturbances. Even the thoughts, as measured by the so-called 
“ association ” tests and the finger prints are marvellously similar. The 
dissimilarity of environment has had little effect on altering the rhythm of 
development, which is controlled by an internal mechanism. The two-egg 
twins are merely ordinary brothers and sisters who are born simultaneously, 
and though the intrauterine environment and that of early years is as nearly 
identical as possible, yet they are as dissimilar as brothers and sisters are 
apt to be. 

Though human heredity is the leading branch of eugenical research, yet 
it is only one. A fascinating branch of the subject is that of mate selection, 
including a study of those external and internal conditions that control in 
this phenomenon. While propinquity is often considered the all-sufficient 
basis of mate selection, yet statistical research reveals such facts as these; 
that there is a selection of mates of corresponding divergence from the 
mean in stature; that red-haired persons do not marry as frequently as 
expected on a random basis; that persons of opposite temperaments tend 
to marry with each other. 

Research on fecundity, especially the differing fecundity of peoples 
having dissimilar social values in the population has not received the atten¬ 
tion it deserves; still we know something of the fractions of sons and 
daughters of college men and women and have some facts available towards 
a study of fecundity of the socially inadequate. Always, however, it is 
not to be forgotten that it is the residuum of surviving children of a marriage 
that counts in the race and the children of the less socially adequate strains, 
are permitted a larger selective death rate than are those of the more effi¬ 
cient strains. That is one reason why from the less developed strains, 
vigorous and effective progeny are occasionally arising; while some lines 
of the more effective and prosperous families end in weak and lethal descend¬ 
ants. Modern surgery has done much to keep alive weak and defective 
individuals, but little to improve racial qualities. Selection and its effects, 
including those of war, have been all too little studied. 

But fecundity of stocks is only a part of the problem in a country which 
like ours, has in a single year, added about as much to the population by 
immigration as by birth. Probably never before in the world has such a 
migration of all sorts of races in such numbers, over so great a distance, 
taken place. Here in America we have watched the process with mis¬ 
givings, and felt a lack of sufficient knowledge to direct our action. The 
present policy of selecting immigrants is a reasonable one, certainly; and 


RESEARCH IN EUGENICS 


27 


every one who recognizes the effect of quality of the germ plasm on national 
life, hopes it will be continued and extended until we know something of the 
family, as well as individual performance, of each applicant for entry into 
the United States. The best, as well as the most recent study of the effect 
of a mixture of races upon a country is Mr. Charles W. Gould’s “ America: 
A Family Matter,” and his conclusions are not encouraging. But the 
student of human genetics hopes to put this marvellous mixture of races 
to account in his study of human inheritance. The greatest opportunity 
in the world is offered for the study, since nearly all the races of mankind 
can be found in New York City alone, in considerable numbers, talking the 
one language and making mixed marriages, which are often strikingly 
diverse. This is a field that is extremely alluring and which has been 
little worked. 

But I fear I tire you with this prolonged discussion of the results and 
the future of eugenical research. No doubt there are many who are 
inquiring, “But where does environment come in?” And there are others 
who would urge that the great problem for investigation is that of the 
relative importance of heredity and environment. It seems to me that 
we should not formulate the problem in this manner. There is no 
heredity without environment and few environmental effects which are 
not dependent also upon heredity. Schooling is good for those who are 
not feeble-minded; moral training yields excellent results in the case of 
such as have normal inhibitions; musical education is valuable if the 
elements of musical capacity are present; painting lessons are fine if the 
pupil be not color blind. Certainly every child deserves the greatest 
possible opportunities; but the same conditions will be an opportunity to 
him who is able to take advantage of them, and no opportunity to him 
whose hereditary limitations do not enable him to use them. 

And finally, what are some of the practical applications that we may 
expect to be made of eugenical research? One, certainly, is a higher estima¬ 
tion of the importance of hereditary capacities in human behavior. This 
may save us from disregard of innate differences in capacities which lead us 
on the one hand to adjudge all men equally capable of acting in accordance 
with the mores; and, on the other, to explain all offences as due to poor 
environment. Both false views neglect the fact of differences in inborn 
capacities. . 

Again, there will come a realization of the importance of heredity in 
marriage matings. Young persons to whom marriage is so serious a 
matter, will be led to stop and consider, when they feel they are falling in 
love, and inquire concerning consequences to offspring. Already there is 


28 


CHARLES BENEDICT DAVENPORT 


being developed a well defined conscience in the matters of cousin marriages 
and of matings into families with grossly defective members. This is 
shown by the extensive correspondence that the Eugenics Record Office 
has been obliged to enter into with persons who are contemplating marriage. 
They are quite willing to submit an extensive account of their family 
traits; and they write to learn what is known about the inheritance of some 
family weakness or defect. The people who make these inquiries are often 
unusually intelligent and not at all radical; some of them stand high in the 
social world. It is a high idealism and a forward looking one which leads 
them to seek the desired knowledge and one can only respond to these 
requests, telling what is known, or highly probable, in respect to the re¬ 
currence. of the family defects in the offspring. Whether the conclusions 
that one is able to give are always very valuable or not, at least the custom 
of considering children and their inheritance of familial traits is one to be 
encouraged. Normal persons marry to beget normal children and it is 
natural for them to seek information concerning heredity of particular 
traits. 

And again, it may be hoped that the study of racial characters will lead 
men to a broader vision of the human race and the fact that its fate is 
controllable. We may hope that reasonable persons will consider the 
progress of mankind, not by the years or generations merely, but by cen¬ 
turies or millenia. We may learn by the history of mankind in the last 
20,000 years how near it has come to extinction; and we must recognize 
that it will take only a little interference with natural instincts and a little 
interference with natural selection during a few generations to bring the 
species, or one race of it, rather abruptly to an end, just as other human 
races have come to an end in historical times. The human species must 
eventually go the way of all species of which we have a paleontological 
record; already there are clear signs of a wide-spread deterioration in this 
most complex and unstable of all animal types. A failure to be influenced 
by the findings of the students of eugenics or a continuance in our present 
fatuous belief in the potency of money to cure racial evils will hasten the 
end. But if there be a serious support of research in eugenics and a willing¬ 
ness to be guided by clearly established facts in this field, the end of our 
species may long be postponed and the race may be brought to higher 
levels of racial health, happiness and effectiveness. 


GENETIQUE ET ADAPTATION 
L. CUENOT 

University of Nancy , Nancy ? France 

Les recherches genetiques modernes, parmi lesquelles les travaux 
americains tiennent une place preponderate, nous ont appris que le 
potentiel ou patrimoine hereditaire, substratum materiel de ce qui passe 
des parents aux descendants, est constitue d’une part par le cytoplasme 
complique de l’oeuf, d’autre part par les substances nucleates male et 
femelle. La polarite, la symetrie, le type de segmentation et le plan ou la 
position et la proportion relatives des organes futurs sont determines 
uniquement par le cytoplasme de l’oeuf, dans lequel il y a frequemment 
des substances organo-formatrices visibles ou morphoplasmes , mais les 
differentiations des stades ulterieurs du developpement, c’est a dire les 
caracteres specifiques et individuels, sont influences a pouvoir 6gal par 
l’oeuf et le spermatozoide, done par les substances nucleates qui constituent 
l’appareil chromosomien. L’analyse mendelienne revele dans celles-ci 
l’existence de facteurs independants, que l’on peut regarder comme de 
tres petites masses materielles de substances chimiques definies, susceptibles 
de changements ou mutations qui conditionneront un changement somatique 
dans une generation suivante, le changement somatique pouvant 6tre 
extremement minime ou au contraire constituer d’un coup un ecart con¬ 
siderable du type normal. Par extension du terme, comme nous ne nous 
apercevons de la mutation chromosomique que par l’effet qu’elle produit 
sur le soma, nous pouvons appeler aussi mutation la variation heritable. 
On ne sait pas provoquer experimentalement des mutation determinees; 
elles apparaissent isolement, d’une fafon imprevisible; cependant il est 
permis de croire qu’elles sont en relation lointaine avec des changements 
du milieu exterieur (excessive variability des animaux domestiques 
comparee a l’uniformite de leurs souches sauvages, mutations innombrables 
des Achatinelles des lies Sandwich qui n’ont du recevoir cependant a 
l’origine que tres peu d’exemplaires, etc.). On comprend qu’ une modifica¬ 
tion du chimisme general puisse etre produite par un changement de 
climat, par l’introduction dans la nourriture d’une substance nouvelle, et 
que cette modification retentisse sur les cellules du corps, y compris les 
sexuelles; ces dernieres presenteront, a echeance plus ou moins yioign^e, 

29 


30 


L. CUENOT 


des mutations visibles, qui pourront etre la derniere etape de petits change- 
ments invisibles, cumulatifs. Les faits semblent montrer, contrairement 
a ce que croyait Delage, que les individus d’une meme espece different 
assez les uns des autres (bien que nous ayons une tendance a les croire 
identiques) pour que la modification possible les touche tres inegalement, 
et qu’il n’y ait qu’un nombre restreint d’entre eux qui presentent des 
mutations. 

La mutation peut se produire aussi bein dans des cellules sexuelles que 
dans des cellules somatiques initiales de bourgeons quelconques (mutations 
gemmaires); les graines portees par des plantes issues de mutations gem- 
maires peuvent reproduire par voie sexuee le type nouveau, qui peut etre 
egalement propage par bouturage (Pomme de terre). 

On ne voit pas qu’on puisse faire d’objections graves a ces conceptions 
factorielles ou mendeliennes et a celle de la mutation, etayees, semble-t-il, 
sur des preuves sans nombre. Partant de cette base, on peut se demander 
quelle est la position des mendeliens ou mutationistes a l’egard des grands 
problemes de l’heredite des caracteres acquis, de l’origine des adaptations 
et des orthogeneses? Je laisserai de cote le troisieme probleme, dont la 
solution depend beaucoup, du reste, de celles donnees aux deux premiers. 

Le caractere acquis est une modification apparaissant chez un individu, 
plante ou animal, a un age quelconque, qui est visiblement Peffet d’une 
cause exterieure et accidentelle, a tel point que si cette cause n’etait pas 
intervenue, la modification ne se serait assurement pas produite. II est 
bien entendu que le caractere acquis est la difference entre l’etat normal 
ou l’etat qui a servi de point de depart, avant Paction de la cause modifiante, 
et l’etat nouveau, apres action de cette cause. Pour qu’un caractere soit 
considere comme acquis, il faut que la relation de cause a effet apparaisse 
avec evidence, soit qu’on ait fait agir la cause experimentalement, soit que 
l’observation naturelle ait, ce qui est rare, la rigueur et la certitude 
d’une experience. II est inutile d’insister sur l’importance capital du 
phenomene pour l’explication generale de Involution et en particulier des 
adaptations; en effet, comme on pouvait—s’y attendre, les reactions de 
l’individu a une action exterieure quelconque ont tres generalement une 
value adaptative ou protectrice: une peau blanche qui se pigmente au 
soleil protege efficacement contre Paction nocive des ultra-violets, l’im- 
munite garantit contre une nouvelle atteinte de la maladie qui la 
determinee; un muscle, une articulation, exerces dans de certaines limites, 
fonctionnent plus efficacement et plus aisement qu’avant l’entrainement; 
un acte appris dont on a l’habitude passe presque a l’etat de reflexe ou 
d’instinct. Si l’heredite des caracteres acquis existe, si faible soit-elle nous 


GENETIQUE ET ADAPTATION 


31 


possedons la clef d’une quantite enorme d’adaptations; si elle n’existe pas, 
il fauU:rouver d’autres explications tres differentes. Ce n’est pas seulement 
une question qui interesse des savants speculates, renfermes dans leur 
tour d’ivoire; c’est aussi une question d’ordre social. Quand on dit au 
grand public que la pratique des sports, meme excessifs, prepare des genera¬ 
tions nouvelles vigoureuses, on a certainement l’idee que les “athletes 
complets” ou meme differencies par des exercices extraordinaires legueront 
a leurs descendants au moins un rudiment de leurs qualites acquises. C’est 
certainement l’opinion des eleveurs qui pensent que les effets de l’entraine- 
ment des Chevaux de course, de la bonne ou mauvaise nutrition du betail, 
sont transmissibles dans une certaine mesure. 

II y a des categories distinctes de caracteres acquis relevant de causes 
differentes: 1. les mutilations; 2. les effets des maladies parasitaires 
produisant une intoxication generale; 3. Faction des grands facteurs 
naturels, lumiere, temperature, humidite, salure, nouriture; 4. les effets de 
Fusage et ceux du non-usage; 5. les acquisitions du dressage, de l’educa- 
tion intellectuelle ou sportive. II faut reconnaitre que la demonstration 
negative, c’est a dire celle de la nonheredite des caracteres acquis de Fune 
des categories, vaut pour celle-la seulement, et ne saurait legitimement 
etre etendue aux autres: ce sont des problemes distincts. Mais, par contre, 
s’il y avait une experience qui montrait indiscutablement l’heredite d’un 
caractere vraiment acquis, ce serait une probability bien forte pour d’autres 
categories, car bien que Fon ne comprenne absolument pas comment une 
modification corporelle acquise pourrait s’inscrire sous sa forme plus ou 
moins attenuee dans le patrimoine hereditaire, si le fait etait prouve une 
fois, l’argument de Fincomprehension perdrait toute sa valeur. 

Pour la l re categorie de caracteres acquis, on peut dire que la reponse 
est definitive; depuis les critiques et les experiences de Weismann, maintes 
fois repetees, personne ne croit plus a l’heredite des mutilations; les observa¬ 
tions de tous les jours confirment celles des biologistes, et il est certain que 
les pseudo-exemples de transmission de mutilations que Fon rapporte 
assez souvent chez des animaux domestiques et de famille humaine sont 
ou de simple coincidences ou des erreurs d’observation. Les experiences 
de Brown-Sequard, du reste mal faites, qui concernent l’heredite de mutila¬ 
tions et celle de desordres physiologiques consecutifs a des mutilations 
nerveuses, ont ete completement controuvees par des travaux dans lesquels 
on peut avoir confiance (Wrzosek et Maciesza) et il n’en reste rien. 

Divers auteurs ont affirme l’heredite des acquisitions de la 5 m categorie, 
notamment du dressage, quelque peu vraisemblable que cela paraisse; 


32 


L. CUENOT 


en particulier, Hachet-Souplet 1 (1912), rapporte les exemples suivants qu’il 
considere comme demonstratifs: un Singe macague auquel il avait appris, 
non sans peine, a tuer des Rats, donna naissance a des petits qui chassaient 
merveilleusement les Rats; des Chats habitues a respecter les Souris ont eu 
des petits qui ne prenaient pas de Souris, meme quand on retardait inten- 
tionellement la distribution de leur nourriture; des Passereaux habitues 
a tirer la chaine d’un petit puits pendant six generations ont donne naissance 
a des jeunes qui, sans dressage, savaient tirer la m£me chaine. Une chienne 
avait ete dressee a faire des pirouettes rapides a gauche; une fille de cette 
chienne, elevee a la campagne, n’ayant aucun exemple sous les yeux et 
n’ayant regu aucun dressage, se mit a faire toute seule des pirouettes a 
gauche vers cinq ou six mois. Tout cela est bien etonnant, et il est permis 
de rester sceptique; bien que l’on ne puisse critiquer des experiences que 
Ton n’a pas suivies, je suis persuade qu’il y a un “trou” dans ces observa¬ 
tions, du peut-etre a la supercherie des aides et a des dressages subreptices 
demeures inapergus, et je ne doute pas que l’heredite des habitudes aille 
rejoindre celle des mutilations. C’est dire que nous ne croyons nullement 
que les instincts sont des habitudes acquises, devenues hereditaires. 

La 3 me categorie concerne les facteurs de milieu; personne ne doute que 
les facteurs de milieu ont une influence determinante extraordinaire sur 
les caracteres des animaux et des plantes, et que, lorsqu’on les change 
experimentalement jusqu’aux limites extremes compatibles avec la vie, 
il en resulte souvent des modifications notables chez les individus soumis 
des modifications notables chez les individus soumis des le jeune age au 
changement d’ambiance. Mais la question est de savoir si ces modifications 
passent, meme a un degre tres diminue, a la generation suivante elevee en 
milieu normal; si oui, nous comprenons facilement la formation des races 
ou sous-especes geographiques; en effet, les resultats de l’action de milieu 
seraient necessairement cumulates, et apres un nombre suffisant de genera¬ 
tions passees dans le milieu modifiant, l’espece pourrait etre tres notable- 
ment transformee, le plus souvent dans un sens utile (car, sans cela, elle 
ne resisterait pas au changement), et, m£me en revenant au milieu normal, 
l’espece nouvelle pourrait conserver ses caracteres acquis. Si non, les effets 
de Paction de milieu se produisent a nouveau pour chaque individu, il n’y 
a pas de modifications cumulatives, et l’influence de l’ambiance sur le 
corps n’a aucun interet au point de vue de Involution. 

On a fait beaucoup d’experiences sur les facteurs de milieu, surtout avec 
des Insectes: des centaines d’essais ont donne des resultats negatifs au point 

1 Hachet-Souplet, La genese des instincts, etude experimentale (Biblioth. de Philos, 
scient., Flammarion, Paris, 1912). Voir p. 239. 


GENETIQUE ET ADAPTATION 


33 


de vue de la transmission du caractere acquis; quelques-uns seulement 
(Standfuss, E. Fischer, Schroder) ont eu un resultat faiblement positif, 
mais ils sont insuffisamment probants, les auteurs ayant opere avec des 
especes naturellement si variables qu’on est en droit de se demander si le 
caractere de coloration qu’ils ont cru acquis n’existait pas avant l’experience 
chez certains individus. Les experiences de Kammerer sont plus embar- 
rassantes encore: il a publie de 1904 a 1911 un grand nombre de travaux 
demontrant la considerable influence de l’ambiance sur les modes de re¬ 
production des Amphibiens (Alytes obstetricans , Salamandra maculosa et 
atra ), sur la couleur d’Amphibiens ( Salamandra maculosa) et de Reptiles 
(Lacerta ), et mettant en evidence dans la plupart des cas la transmission 
hereditaire des caracteres acquis. Ce n’est pas trop dire qu’a premiere 
vue les resultats de Kammerer paraissent incroyables; il disposait, il est 
vrai, d’une installation exceptionnelle, au Prater, a Vienne (Biologische 
Versuchsanstalt de l’Academie des Sciences), mais qu’il ait pu realiser, 
meme avec cette installation, des elevages d’une difficulty aussi grande et 
d’aussi longue duree, est deja surprenant; des le premier jour, les experiences 
de Kammerer ont paru trop bien reussies, trop demonstratives et trop 
extraordinaires pour ne pas cacher des erreurs. Boulenger et Bateson les 
ont critiquees severement; on a parle meme de tricheries et de substitution 
de preparations; Hans Przibram, qui dirige la station du Prater, a bien 
voulu me dire qu’il n’en etait rien, maisc’estaumoins une indication que les 
resultats de Kammerer ont rencontre generalement l’incredulite./ Si 
Przibram ne les avait pas patronnes, pour ainsi dire, en rendant compte 
des travaux de Kammerer dans son excellent livre Experimental-Zoologie 
(tome 3, 1910), j’imagine qu’on aurait attendu, pour en tirer argument, 
qu’ils aient ete confirmes par un ou deux observateurs d’autres pays, ce 
qui n’a pas encore eu lieu, tout au contraire. 

En somme, jusqu’ici l’experience decisive, verifiee certaine manquait 
encore, et les biologistes avaient le droit de ne pas croire a l’heredite des 
caracteres acquis de la 3 me categoric (facteurs de milieu) et de la 2 me (im¬ 
munity), mais le probleme restait toujours pose; cela est si vrai, que la 
croyance a l’heredite de ces caracteres acquis etait presque une affaire 
de foi, de nationality; la grande majority des zoologistes frangais tenait pour 
l’affirmative, a la fuite de Giard, d’Edmond Perrier, de Le Dantec, de F. 
Houssay, de Delage, tous plus ou moins lamarckistes, les Americains, sauf 
quelques paleontologistes, penchaient plutot pour la negative. Mais la 
question a pris recemment une face nouvelle avec les experiences de 
M. F. Guyer et E. A. Smith (1920). 2 

» 

2 Guyer et Smith, Studies on cytolysins. 11. Transmission of induced eye-defect 
(Joum. of exp. Zool., 31, 1920, p. 171). 


34 


L. CUENOT 


En injectant a des Poules des cristallins broyes de Lapins, ils obtiennent 
un serum cristallolytique; celui-ci est injecte a des Lapines albinos pleines 
depuis 10 a 13 jours (epoque particulierement importante pour le developpe- 
ment du cristallin); la cristallolysine de la Poule n’agit pas sur les cris¬ 
tallins des Lapines adultes, peut-etre parce qu’elle n’atteint pas ces organes, 
alors depourvus de vaisseaux, mais elle passe a travers les placentas, et affecte 
plus ou moins fortement les foetus infermes dans l’uterus; beaucoup meu- 
rent; ceux qui survivent presen tent parfois (9 cas sur 61 petits) un cristallin 
plus petit et plus ou moins opaque, ou bien de la microphtalmie ou meme 
une fonte complete de l’oeil. Sans aucun doute, cela est bien du a Paction 
specifique de la cristallolysine, car les Lapins de controle (48), provenant 
de meres injectees soit avec du serum de Poules normales, soit avec du 
serum de Poules preparees avec un tissu de Lapin autre que le cristallin, 
n’ont presente aucune modification de l’appareil visuel. Ce n’est pas non 
plus une coincidence fortuite due a une mutation passee inapergue, car on 
ne connait dans aucune race de Lapins d’animaux a yeux naturallement 
defectueux. Jusqu’ici il n’y a rien d’absolument nouveau ni de surprenant, 
car on savait deja que des serums cytotoxiques ou des anticorps immuni- 
sants peuvent passer a travers les placentas pour aller impressionner des 
organes de foetus. Bien entendu, les yeux defectueux des jeunes constituent 
un caractere acquis certain, car si l’on n’avait pas injecte a leurs meres de 
serum cristallolytique, les yeux seraient restes sains. 

Or, ce caractere acquis s’est montre hereditaire] il a ete transmis jusqu’a 
la 8 me generation sans autre traitement que les injections originelles; 
Panomalie tend a devenir plus grave dans les generations successives et 
aussi a apparaitre dans un nombre plus grand de jeunes; la transmission est 
souvent d’un type irregulier, tantot l’oeil droit seulement ou le gauche, 
tan tot les deux yeux etant atteints; mais dans les dernieres generations, 
sans doute parce que les parents sont pris de preference parmi les individus 
les plus touches, il y a un nombre croissant de jeunes qui ont les deux yeux 
affectes. Pour demontrer que le reapparition de la dystrophie specifique 
est bien due a l’heredite et non a un passage d’anticorps une fois fabriques, 
Guyer et Smith ont croise des males defectueux avec des femelles normales 
(sans relation familiale avec les stocks traites): la l re generation a des 
yeux constamment normaux (dominance du caractere yeux normaux), 
mais les femelles heterozygotes de cette generation, croisees a nouveau 
avec des males defectueux, ont donne un certain nombre de petits a yeux 
degeneres. Au point de vue genetique, cela ne peut se comprendre que 
si Pon admet que le patrimoine hereditaire des males a subi une modification 
specifique. 


GENETIQUE ET ADAPTATION 


35 


Bien entendu, on ne pourra faire fond sur l’experience de Guyer et Smith 
que le jour ou elle aura ete refaite, avec les memes resultats, par d’autres 
observateurs; mais il semble bien que les deux auteurs ont opere avec soin, 
et ont prevu et evite les causes d’erreur et les critiques possibles. Si l’ex- 
perience de Guyer et Smith est bonne et valable, que de consequences graves 
et que changements dans nos manieres de voir! 

Puisque des anticorps specifiques peuvent produire des modifications 
specifiques dans les cellules germinales, nous ne pouvons pas echapper a 
cette consequence qu’ il y a dans le patrimoine hereditaire une substance 
ou une particule qui a une certaine correspondance avec le cristallin de- 
veloppe, puisque la cristallolysine (qu’elle soit substance ou propriete, 
peu importe) touche d’une part les cristallins en voie de formation du foetus, 
et d’autre part les cellules germinales de celui-ci, de telle sorte que ces 
dernieres seront l’origine de petits a cristallin defectueux. On comprendrait 
a la rigueur que la cristallolysine, a la fa$on d’un poison, modifie quelque 
peu la cellule germinale, mais d’une fa^on quelconque, de maniere a ce qu elle 
donne naissance a un petit plus ou moins dystrophie; mais que la modifica¬ 
tion soit specifique, voila qui est stupefiant! Il y a done dans le germe une 
partie qui est “ representative ” du cristallin, par exemple un colloide de¬ 
termine analogue aux colloides du cristallin developpe. Et s’il y en a pour 
le cristallin, il doit y en avoir pour toutes les cellules differenciees de l’organ- 
isme, les pigmentaires, les nerveuses, les adipeures, les stomacales, les 
renales, etc. Nous revenons ainsi a une theorie particulaire de la constitu¬ 
tion des cellules germinales, a des determinants chimiques, theorie que Delage 3 
a autrefois formulee d’une fagon tres nette (1895), en reprenant et modifiant 
des idees plus ou moins analogues de Weismann et de W. Roux. Le germe 
serait un microcosme renfermant tons les colloides difierents de Vorganisme 
aduite; si l’experience de Guyer et Smith est vraie, je ne vois pas qu’on puisse 
echapper a cette consequence. Elle n’a du reste rien de contradictoire 
avec la conception mendelienne, car le determinant chimique ou colloide 
representatif parait bien se confondre avec le facteur mendelien. En 
effet, Guyer et Smith montrent que l’element germinal touche par la cr i s_ 
tallolysine se comporte a peu pres comme un facteur mendelien recessif. 
quand des males ou des femelles a yeux anormaux sont croises avec des 
individus non traites, provenant pour plus de surete d’autres regions, tou- 
jours la progeniture apparait avec des yeux normaux; mais les petits sont 
des heterozygotes et renferment a l’etat domine le caractere yeux defectueux ] 
en effet, si ces Lapins d’apparence normale sont croises avec d autres a 

8 Delage, La structure du protoplasma et les theories sur l’heredite, etc. Reinwald, 
Paris, 1895 (voir p. 807). 


36 


L. CUENOT 


yeux defectueux, il apparait cette fois dans leur descendance un nombre 
plus ou moms grand de petits a yeux anormaux. Deux individus a yeux 
defectueux (homozygotes domines), croises ensemble, devraient, dans 
l’hypothese de facteurs mendeliens simples et typiques, donner uniquement 
des petits presentant Panomalie; il n’en est pas toujours aussi, mais il y a 
une majorite frappante de cristallins opaques ou petits dans leur descend¬ 
ance. Les documents sont encore insuffisants pour voir clair dans la 
gehetique du caract&re, mais en gros, abstraction faite de details qui s’eclair- 
ceront probablement plus tard, les caracteres yeux normaux et yeux lyses 
forment en couple allelomorphique, avec dominance du premier. 

Est-ce a dire que l’experience de Guyei et Smith, meme si elle etait 
confirmee dans tous ses details est parfaitement satisfaisante? Il s’en 
faut de beaucoup; un mauvais sort veut que les preuves de l’heredite des 
caracteres acquis ne presentent jamais cette plenitude d’evidence qui 
entraine irresistiblement la conviction et laisse Pesprit en repos. La 
cristallolysine a une action vraiment deconcertante: elle n’agit pas sur les 
cristallins de la mere gestante, premiere anomalie qui, il est vrai, a regu 
une explication plus ou moins bonne; elle agit capricieusement sur les cristal¬ 
lins en voie de developpement des foetus (9 fois sur 61) et d’apres l’hy- 
pothese, sur les determinants cristalliniens des cellules germinales de ces 
memes foetus. On doit done s’attendre a ce qu’elle touche egalement les 
cellules germinales dans les ovaires de la mere gestante, aussi bien que 
celles des foetus; or, il n’en est rien: les Lapines qui survivent aux injections 
de serum anticristallin et qui ont donne des jeunes a yeux anormaux, sont 
couvertes a plusieurs reprises par les memes males normaux, apres que 
les traitements seriques ont ete arretes (depuis combien de temps?), et pas une 
trace de malformation oculaire n’est visible dans leur descendance, cepen- 
dant nombreuse. Il y a la quelque chose de tout a fait incomprehensible, 
a moins d’admettre que les cellules germinales ne peuvent etre modifiees 
qu’a des stades particuliers, celui de la periode de multiplication des gonies, 
ou bien celui de la synapse. Or, on sait que chez les femelles de Mammi- 
feres, ces phenomenes se passent chez Pembryon, et que l’adulte n’a plus 
que des ovocytes qui s’accroissent lentement et inegalement pour arriver 
successivement a maturite. Nous avons un moyen de verifier cette hypo- 
these subsidiaire, car les males adultes presentent toutes les phases de la 
spermatozoide; an injectant du serum anticristallin a des males, on devrait 
toucher leurs cellules sexuelles a la periode sensible; et accouples plus tard 
a des femelles normales, ils devraient transmettre Panomalie, qui ne serait 
visible que chez leurs petits-enfants. -C’est une experience cruciale a faire; 
je ne vois pas que Guyer et Smith Paient tentee. Il sera indique, pour 


GENETIQUE ET ADAPTATION 


37 


simplifier l’experience, de ne pas utilise seulement la Poule comme source de 
cristallotoxine, mais bon le Lapin lui-meme, pour ne pas introduire le f acteur 
etranger serum de Poule. Du reste Guyer 4 a reussi recemment (1921) 
& obtenir un jeune Lapin a deux yeux defectueux en injectant a plusieurs 
reprises a sa mere normale des cristallins broyes de Lapins, avant qu’elle 
fut pleine et pendant sa gestation. Un Lapin peut done fabriquer un 
anticristallin—Lapin aussi efficace que celui engendre par une espece 
etrangere. 

Revenons aux caracteres acquis: les experiences de Guyer et Smith, en 
les supposant confirmees dans le fait et dans Interpretation, permettent 
d£sormais d’accepter l’heredite des caracteres acquis sous l’influence 
des grands facteurs generaux du milieu; en meme temps que ceux-ci 
produisent leur effet plus ou moins adaptatif sur certains tissus du corps, 
ils modifient d’une fayon parallele les colloides representatifs des cellules 
germinales, de telle sorte qu’il pourra y avoir effet cumulatif du meme 
facteur general agissant sur des generations successives. C’est exacte- 
ment ce qui a ete suppose depuis longtemps par Weismann luimeme, et 
ce que l’on a designe sous le nom d 'induction parallele (mot de Detto). 
Weismann a illustre le fait par l’exemple celebre du Chrysophanus phlaeas: 
ce Papillon, a tres large repartition geographique, varie comme beaucoup 
d’autres suivant les saisons et les localites: dans la region septentrionale, 
l’aile superieure est rouge dore avec bordure noire et points noirs sur le 
disque, l’aile inferieure noiratre avec une bande submarginale rougeatre; 
dans l’Europe meridionale, on trouve dans la generation d’ete des 
exemplaires plus grands, dont les deux ailes sont presque entierement 
noires (forme eleus), avec tous les intermediates entre cette forme et 
le type. Si Pon eleve en Allemagne des chenilles provenant de phlaeas 
de Naples et qu’on soumette les pupes a une basse temperature (10°), il 
se developpe des Papillons un peu moins noirs que ceux qui volent a Naples, 
mais beaucoup plus que les allemands; au contraire, des pupes d’origine 
allemande soumises a une temperature elevee (38°) donnent des Papillons 
qui sont un peu moins rouge feu et un peu plus noirs que les Papillons alle¬ 
mands habituels. Le caractere geographique et hereditaire des races 
meridionales est done dirige dans le meme sens que Paction d’une tempera¬ 
ture elevee sur les pupes; et il parait vraisemblable de supposen que lorsque 
l’espece, primitivement septentrionale, a gagne les regions du midi, l’eleva- 
tion de temperature, agissant pendant de nombreuses generations, a 
affecte avec un resultat parallele les elements formateurs de couleur des 
ailes et les colloides representatifs du pigment dans les cellules germinales. 

4 Guyer, Immune sera and certain biological problems (Amer. Natur., 55, 1921, p. 97). 


38 


L. CUENOT 


Jusqu’ici on pensait que l’immunite, caractere acquis apparu apres une 
maladie microbienne, n’etait pas transmissible, si ce n’est a une premiere 
generation par echange placentaire; or Guyer (1921) rapporte que des 
Lapins injectes successivement avec le vaccin typhique et des germes 
typhiques vivants peuvent transmettre a leurs jeunes et meme a la genera¬ 
tion suivante la propriete d’agglutiner des bacilles typhiques dans du serum 
dilue. Si cette experience, en contradiction avec les travaux anterieurs, 
est confirmee, on pourra comprendre pourquoi les habitants d’une contree 
oil sevit une maladie habituelle sont en general plus aptes a y resister que 
ceux d’une contree indemne, la maladie etant moins severe dans le premier 
milieu, ou bien le nombre des individus absolument immune y etant plus 
considerable. II y aurait, en meme temps qu’une selection des plus 
resistants, transmission d’une immunite partielle. 

Reste la categorie la plus importante, les caracteres acquis par 1’usage 5 et le 
non-usage, question qui creuse le fosse profond entre les lamarckistes et 
les opposants. La question n’est pas abordable experimentalement; 
aucune experience, meme prolongee pendant plusieurs generations humaines, 
ne montre ni ne peut montrer l’heredite des effets de l’adaptation fonction- 
nelle individuelle; aussi les lamarckistes ont-ils cherche la demonstration 
dans des observations naturelles; on connait en effet un certain nombre de 
caracteres hereditaires, apparaissant deja chez les embryons en dehors de 
toute cause exterieure, qui semblent resulter de la fixation de reactions 
individuelles a la pression, a l’exercice ou au defaut d’usage; ce ne peut 
etre qu’un argument de vraisemblance, mais une accumulation d’exemples 
de cette sorte pourrait toutefois avois une valeur demonstrative, bien que 
l’on ne comprenne absolument pas comment un effet mecanique, produit 
chez l’animal developpe, pourrait s’inscrire dans le patrimoine hdrdditaire. 
Les meilleurs exemples lamarckistes sont: l e la ressemblance des courbures 
du corps d’un Poisson bon nageur avec celles d’un modele plastique soumis 
aux memes courants d’eau qu’un Poisson en nage filee (F. Houssay); 2® 
la perforation de la paroi operculaire des tetards de Bombinator , meme en 
l’absence des pattes anterieures qui provoquent normalement cette 
perforation (Braus); 3 e l’echancrure de la levre superieure du Sanglier male, 
en rapport avec la canine relevee en dehors; 4 e les callosites des surfaces 
plantaires des animaux marcheurs (Semon); 5 e les callosites des surfaces 
(autres que les plantaires), sur lesquelles s’appuient divers animaux, comme 
les Chameaux, la Girafe, le Phacochere, les Autruches (Leche, Duerden, 
etc.); 6 e l’atrophie des yeux chez les animaux cavernicoles ou abyssaux. 

6 Par le mot usage, on entend non-seulement 1’usage proprement dit, mais aussi tous 
les effets m6caniques de traction, pression, etc. 


GENETIQUE ET ADAPTATION 


39 


Je n’ai pas l’intention de critiquer en detail ces observations, dontbeaucoup 
meriteraient d’etre reprises et etudiees tres en detail; je me bornerai a 
quelques remarques. Un Sus scrofa male adulte presente comme on sait 
des canines superieures fortement relevees, qui retroussent la levre; on a 
evidemment l’impression que c’est la pression exercee par la dent, objet dur, 
sur les parties molles, qui conditionne cet etat; or, si on examine un embryon 
male, avant la poussee des canines, n constate (fig. 1) que la levre superieure 
presente deja une echancrure manifeste, a l’endroit ou la dent passera bien 
plus tard. Mais on peut formuler une toute autre hypothese que celle 
de l’heredite d’un caractere mecaniquement acquis; il est evident qu’il 



Fig. 1. Tete de foetus de Sanglier (Sus scrofa L.) montrant en X le pli preforme de la 
levre superieure. Gr. nat. 

y a chez le male une cause interne, dispositif osseux ou autre, qui contramit 
la canine a pousser en dehors et en haut, au lieu de croitre vers le bas: or, 
n’est-ce pas ce dispositif qui, deja chez l’embryon, tira sur la levre superieure 
et y determine la petite echancrure? 

Les callosites des surfaces d’appui,rexemplede beaucoup le plus favorable 
aux lamarckistes, sont evidemment determinees chez l’embryon par une 
cause actuelle, probablement la pression de surfaces osseuses sur la region 
epidermique; si l’animal developpe s’appuie sur ces callosites, comme 
l’Autruche etudiee par Duerden, est-ce leur presence, anterieure a l’usage, 
qui a permis a l’animal d’adopter son attitude speciale, ou bien l’animal 
a-t-il pris l’attitude avant d’avoir des callosites? L’un n’est pas plus 
certain que l’autre. 



40 


L. CUENOT 


Quant aux atrophies oculaires des animaux vivant dans l’obscurite 
absolue, il n’est pas impossible, en se basant sur les resultats de Guyer et 
Smith, de comprendre leur genese; des oirganes tels que les yeux, qui cessent 
completement de fonctionner, pourraient produire des modifications humo- 
rales specifiques, des sortes de lysines qui affecteraient les colloides repre- 
sentatifs dans les cellules germinales, et ameneraient graduellement la 
regression hereditaire. 

En resume, il me semble que les mendeliens qui, jusqu’ici niaient com¬ 
pletement l’heredite de tous les caracteres acquis, meme sous la forme de 
l’induction parallele, ne peuvent plus se montrer aussi affirmatifs; il est 
absolument necessaire de verifier ou d’infirmer les experiences de Guyer et 
Smith, en les recommengant avec plus de rigueur,pourmettreenfinun terme 
a la question irritante de l’heredite des acquisitions somatiques et nous 
fixer sur la constitution intime du patrimoine hereditaire. 

Il n’y a pas de mot plus frequemment employe en Biologie que celui 
d 'adaptation, non sans quelque confusion dans l’emploi de ce terme. Tantot 
on entend par adaptation le fait de l’accommodation d’un individu a des 
conditions de vie nouvelles pour lui, c’est a dire les modification somatiques 
plus ou moins favorables, adaptatives comme Ton dit, qui resultent de 
Taction de ces nouvelles conditions; c’est la regulation ou l’adaptation 
fonctionnelle. 

Tantot on appelle adaptation l’accord general des caracteristiques d’une 
espece et des conditions physiques et biologiques du milieu ou elle vit et 
ou elle persiste. Il est evident qu’un animal est toujours adapte aux 
conditions de son milieu: il a un appareil digestif approprie a la nourriture 
qu’il peut se procurer, un systeme locomoteur convenable, une fecondite 
capable d’egaler au moins le taux de destruction normal par carnassiers et 
parasites, des reactions a la lumiere, a la chaleur, etc., qui sont d’accord avec 
sa structure et ses besoins. L’adaptation d’une forme a son milieu, phe- 
nomene global qui ne se constate qu’apres coup, est par definition necessaire 
et suffisante, quels qu’en soientles details. Le jour ou par suite d’un change- 
ment du milieu physique oude l’arrivee de nouveaux concurrents et parasites, 
l’espece n’a plus dans l’ensemble un total qui est en sa faveur, elle emigre 
si elle peut le faire, ou bien elle s’amoindrit et disparait, quelle qu’ait pu 
etre dans le passe la perfection de' ses caracteristiques, tels les Ammonites, 
les Ichthyosaures, les gigantesques Dinosauriens, etc. La selection natu- 
relle est la balance qui pese la somme des caracteres des especes et des 
individus, et qui elimine ceux qui sont trouves trop legers. 


GENETIQUE ET ADAPTATION 


41 


Cette adaptation generale est decoupee quelque peu artificiellement en 
adaptations speciales, en rapport avec des conditions determines du milieu: 
par exemple, on dit couramment que la palmure des pattes des Vertebres 
superieurs est une adaptation a la vie aquatique, parce qu’on voit bien 
l’utilite d’une rame pour le nageur, et que la palmure a apparu independam- 
ment chez beaucoup d’especes aquatiques de groupes differents; mais si 
repandue qu’elle soit, cette adaptation speciale n’a pas un caractere de 
necesessite, puisque la Poule d’eau ( Gallinula chloropus) a les doigts bien 
separes, tout en etant excellente nageuse, que le Foulque (Fulica atra ) 
n’a qu’une frange digitale, et que le Flamant (. Phoenicopterus roseus ) qui ne 
nage que rarement, a une palmure complete! II y a aussi des palmures 
chez des animaux absolument terrestres, soit arboricoles, comme le Galeo- 
pitheque et le Gecko Ptychozoon, Malaisie, soit habitant les dunes, comme 
le Lezard Palmatogecko Rangei (S. O. Afrique). Chez une forme qui a 
change completement de mode de vie, il peut tres bien persister des carac- 
teristiques qui etaient jadis des adaptations speciales a un autre milieu, 
et que l’espece, dans son nouvel entourage, utilise autrement. L’outil 
le plus precieux de l’Homme, sa main prenante, est un reste d’une ancienne 
adaptation a la vie arboricole. 

II est tres digne de remarque que Lamarck et Darwin, de meme que leurs 
successeurs immediats, n’ont nullement cherche a expliquer l’adaptation 
suffisante: leurs theories portent toutes sur un point tres particulier, revolu¬ 
tion des adaptations speciales et des regressions, en somme sur les ortho¬ 
geneses. Pour reprendre les exemples qu’ils ont eux-memes choisis, Darwin, 
etudiant la regression de l’oeil de la Taupe, suppose que la Taupe vit dans 
une galerie souterraine avant que l’oeil ait diminue; s’occupant du planeur 
du Sciuroptere, il regarde son ancetre comme un Ecureuil, done arboricole. 
Lamarck, pour la palmure des aquatiques, part d’un Oiseau a doigts separes, 
mais deja nageur; pour l’allongement des pattes et du cou d’un Echassier, 
il admet un Oiseau de rivage oblige de chercher sa nourriture au fond de 
l’eau. Pour Houssay, l’un des lamarckistes les plus excessifs de l’ecole 
frangaise, le corps du Poisson est le resultat du modelage par l’eau, mais 
il imagine comme point de depart un animal qui vit dans l’eau et qui nage 
vite, done forcement un Poisson. Toutes ces theories prennent done au 
debut l’animal dans un milieu determine, auquel il est evidemment adapte 
d’une fagon suffisante, puisqu’il y vit et y persiste, cette adaptation globale 
comportant necessairement des adaptations speciales. C’est la genese 
de l’adaptation generale que nous allons etudier. 

Reprenant une opinion dont on trouve deja une trace dans Buffon, 
j’admets que l’animal a tient sa destinee des organes avec lesquels il est 


42 


L. CUENOT 


ne, ” c’est a dire qu’il ne peut vivre et durer que s’il a eu la chance de rencon- 
trer un milieu adequat a sa structure et auquel il peut accommoder sa 
physiologie; ne peuvent peupler un milieu que les especes ou les individus 
qui presentent avant d ’y entrer l’adaptation generale necessaire et suffisante, 
qui en un mot y sont preadaptes. C’est aussi l’opinion de De Vries “le 
milieu a seulement choisi les formes aptes parmi la foule et n’a pas de rela¬ 
tion quelconque avec leur origine*” de Th. Morgan “la forme apparait 
independamment du milieu; une fois apparue, elle peut se perpetuer dans 
des conditions convenables, ” de Davenport “la structure existe d’abord 
et l’espece cherche du rencontre le milieu qui repond a sa constitution 
particuliere, ” et d’autres encore. 

L’exemple suivant, que j’ai deja cite ailleurs, 6 permet de bien comprendre 
ce qu’est cette preadaptation generale: il existe en Lorraine des mares ou 
ruisseaux sales, formes par des sources naturelles ayant traverse des depots 
saliferes triasiques ou par des dechets de salines; la concentration en sels 
est tres variable d’une station a une autre, et dans une meme station 
suivant la saison, l’eau a peine saumatre en hiver pouvant devenir sursalee 
au fort de l’ete. Ce milieu est habite par une petite faune riche en individus, 
mais peu nombreuse en especes, qui provient evidemment de l’eau douce 
avoisinante; parmi ces habitants de l’eau salee, je n’en retiendrai qu’un, 
l’Epinoche (Gasterosteus aculeatus), qui s’y trouve en grand nombre et s’y 
reproduit; ce Poisson y atteint une plus grande taille que dans les ruisseaux 
d’eau douce voisins, et presente certaines modifications dans le nombre 
des plaques osseuses qui recouvrent en partie les flancs de l’animal. Cette 
propriete de vivre continuement dans l’eau salee a salure tres variable, 
cette adaptation generale a-t-elle ete acquise par un phenomene lent d’accli- 
matation et de selection? Pas du tout. L’Epinoche est douee d’une 
faculte qui est restee ignoree jusqu’au jour ou on l’a mise en evidence d’une 
fajon experimentale, celle de supporter des changements instantanes de 
salinite; on peut, en effet, transporter brusquement des Epinoches de l’eau 
douce dans de l’eau de mer, et vice versa, sans les tuer; il y a assurement 
quelques individus qui meurent, mais beaucoup survivent indefinement. 
Si on faisait la meme experience avec les autres Poissons d’eau douce, ils 
mourraient certainement au bout de quelques minutes. Ce qui montre 
bien que cette euryhalinite est un phenomene d’ordre cellulaire, lie a une 
resistance speciale des tissus peripheriques a l’osmose, c’est que l’Epinoche 
peut vivre egalement dans de l’eau glycerinee et dans une solution de sucre 
a 10 pour 100 (Siedlecki). A l’etat normal cette propriete est assurement 
indifferente pour l’Epinoche qui habite en eau douce; elle est fortuite, sans 

6 L. Cuenot, Theorie de la preadaptation (Scientia, 16, 1914, p. 60). 


GENETIQUE ET ADAPTATION 


43 


utilite, est un caractere de hasard; mais elle joue un role decisif lorsque 
1’Epinoche a l’occasion d’occuper des eaux saumatres et sursalees, et elle 
prend alors la signification d’une preadaptation. 

II ne manque pas d’exemples analogues: toutes les fois que l’on etudie 
le peuplement d’une place vide actuelle, c’est a dire d’un milieu habitable 
qui se cree en un point donne par suite de circonstances cosmiques ou d’inter- 
ventions humaines (ile volcanique qui surgit des flots, eaux thermales ou 
salines, galeries de mines profondes, conduites d’eau de ville, plantes 
exotiques acclimatees), on constate que des etres vivants ne tardent pas 
a occuper la place vacante; dans tous les cas, le peuplement est opere par 
les animaux ou les plantes du milieu analogue le plus voisin, mais ceux-la 
seulement qui sont capables d’y arriver de par leurs reactions sensorielles, 
de s’habituer a ses conditions speciales et de s’y multiplier. En somme il 
y a filtrage de la faune et de la flore avoisinantes; seules passent a travers 
les mailles du filtre les especes dont l’adaptation generale, convenable pour 
le milieu dont elles sortent, est aussi convenable pour celui ou elles entrent, 
bien que ce dernier puisse etre tres notablement different du precedent. 

Cette seconde adaptation generale peut se decouper aussi en adaptations 
speciales, repondant aux exigences nouvelles; mais ce qui est le plus interes- 
sant, c’est que celles-ci pouvaient etre anterieurement des caracteristiques 
indifferentes ou d’une utilite mediocre, n’ayant pas la valeur de necessites, 
ou des adaptations qui jouaient un role different; j’ai cite plus haut l’eury- 
halinite de l’Epinoche, propriete dont il ne fait assurement pas usage quand 
il vit en eau douce; en void un autre exemple, non moins demonstratif: 
une adaptation frequente des animaux d’eau douce est l’existence chez 
ceux-ci de gros oeufs peu nombreux, a developpement direct donnant nais- 
sance a des petits, capables de mener immediatement le meme genre de vie 
que leurs parents: on connait par exemple les gros oeufs des Ecrevisses 
(. Potamobins , Cambarus, etc.), des Crabes d’eau douce ( Potamon ), des 
Paloemon d’eau douce, du Prosobranche vivipare Paludina, etc. Les 
formes marines, au contraire, on des oeufs petits et nombreux donnant nais- 
sance a des larves nageuses qui menent plus ou moins longtemps la vie 
pelagique, et doivent necessairement subir des metamorphoses compli- 
quees avant d’acquerir la forme definitive de l’espece. On voit facilement 
la raison d’etre de cette adaptation: comme la penetration des animaux 
marins dans l’eau douce s’est surtout operee par les estuaires des grands 
fleuves, on comprend que des larves nageuses auraient ete forcement en- 
trainees au loin par les courants souvent violents des cours d’eau; il est 
done tres avantageux pour des animaux d’eau douce d’avoir des oeufs a 
developpement direct; l’espece peut ainsi remonter graduellement les cour- 


44 


L. CUENOT 


ants et s’etendre en amont, sans risquer d’etre refoulee a chaque genera¬ 
tion. Cela est si vrai que les deux habitants d’eau douce qui, par exception, 
ont des larves pelagiques, la Crevette Atyaephyra Desmaresti et le Mollus- 
que Dreissensia polymorpha, n’ont etendu que recemment leur aire geo- 
graphique, et cela grace a la creation des canaux, dont le courant est trop 
faible pour emporter les larves. Or, on dit sou vent que la presence de gros 
oeufs a ete determinee, apres Ventree en eau douce , par une influence speciale 
de ce milieu; je ne le pense pas; il est beaucoup plus vraisemblable de sup- 
poser que parmi les animaux marins euryhalins qui ont penetre dans les es- 
tuaires, quelques-uns avaient deja de gros oeufs, et que c’est cette preadap¬ 
tation qui leur a permis de franchir la barriere et de s’etablir definitivement 
en eau douce; on connait du reste de gros oeufs a developpement direct 
chez des types franchement marins, comme le Homard et le Nophrops, 
tous deux proches parents de l’Ecrevisse, et chez des Littorines vivipares. 

Les exemples de preadaptations abondent: des animaux littoraux peuvent 
avoir des caracteres de formes abyssales ou pelagiques, des animaux aqua- 
tiques de mer ou d’eau douce presentent des dispositifs propres aux ani¬ 
maux terrestres, de simples lucifuges ont des traits de vrais cavernicoles. 
On peut poser en regie generale que les caracteristiques ou adaptations 
speciales a un milieu donne se retrouvent ga et la, eparses chex les habitants 
du milieu le plus voisin, a l’etat de proprietes latentes ou indifferentes ou 
depassant les necessites. Grace a ces preadaptations, certaines especes 
qui en reunissent un nombre suffisant peuvent quitter leur habitat originel 
pour passer dans un milieu nouveau, inhabite; elles y vivent a l’abri de 
toute concurrence, s’y multiplient; y varient, et peuvent devenir la touche 
de nouveaux groupes differencies. Specialement interessantes sont au- 
jourd’ hui les formes qui restent pour ainsi dire a la limite de deux milieux, 
comme les Tsopodes et Amphipodes littoraux qui ne sont plus marins et 
pas encore terrestres, beaucoup de Batraciens mi-aquatiques mi-terrestres, 
les animaux et les plantes d’estuaires a moitie chemin entre l’habitat marin 
et l’eau douce, les animaux d’ entrees de caves qui hesitent entre la lumiere 
et l’obscurite. II suflira de peu de chose pour que soit realisee l’adaptation 
necessaire et suflisante qui permettra de passer la limite, comme l’ont 
fait autrefois les Cloportes et certains Talitrus et Orchestia terrestres, la 
Dreissensia, les Caridines, Palemons et Crabes d’eau douce, les Araignees 
cavernicoles, etc. 

Les aphorismes suivants resument notre fafon de comprendre l’adapta- 
tion: une adaptation suflisante est necessairement anterieure a l’installa- 
tion dans un milieu donne.—II n’y a done pas de lieu causal entre l’adapta- 
tion suflisante a un milieu et les conditions de ce milieu.—La necessite 


GENETIQUE ET ADAPTATION 


45 


et l’organe creent la fonction.—Ce n’est pas la nourriture qui influence la 
forme des dents, c’est celle-ci qui determine le choix de la nourriture (0. 
Aichel).—Ce n’est pas l’usage de la nourriture cuite qui a amene la reduc¬ 
tion du volume de nos dents et des machoires, si evidente quand on compare 
les premiers Hommes al’Homme civilise actuel; c’est plutot cette regression 
qui nous a impose l’usage d’une nourriture facile a macher.—La Nature 
n’a point taille les dents ihumainesl pour les diverses utilities qu’elles pre¬ 
sented, mais les dents s’etant trouvees, par un arrangement fatal, prendre 
telle ou telle forme, il en est resulte telle ou telle utilite (Philosophes grecs 
predecesseurs d’Aristote).—Ce n’est pas parce que la Girafe broute des 
arbres qu’elle a un long cou, mais c’est parce qu’il lui est venu un grand cou 
qu’elle n’a pu faire autrement que de brouter des arbres.—Les animaux 
cavernicoles entrent dans les cavernes parce qu’ils y trouvent des con¬ 
ditions adequates a leur structure et physiologie. Ils sont cavernicoles 
parce que leurs yeux sont degeneres et parce qu’ils ont perdu leur pigment, 
avant d’etre entres dans les cavernes (Banta).—Les plumes des Oiseaux, 
triple adaptation au vol, a la protection du corps contre la deperdition de 
la chaleur et contre les traumatismes, ont du apparaitre avant ou pendant 
la transformation des membres anterieurs en ailes; je crois qu’on trouvera 
quelque jour un Reptile-Oiseau, anterieur a YArchoeopteryx, ne volant pas 
et ayant deja des plumes.—Les habitats des plantes sont determines par 
leurs particularity de structure et non vice versa (H. B. Guppy). 

II est a peine utile de faire remarques que la notion des preadaptations 
est en parfait accord d’une part avec la conception mendelienne ou factorielle, 
d’autre part avec les idees sur les mutations; celles-ci etant indeterminees 
au point de vue fonctionnel, sans relation utilitaire avec les conditions 
de milieus on ne peut comprendre l’adaptation indeniable des etres a leur 
ambiance qui si elle est le resultat de rencontres heureuses, si l’on vent d’une 
suite d’essais comportant beaucoup d’erreurs et quelques reussites. Nous 
ne connaissons que ces dernieres, aussi tout ce qui survit est necessairement 
adapte au milieu a tous les points de vue, ce qui a donne lieu a Pillusion 
finaliste de l’ecole d’Aristote. 

Si la doctrine genetique moderne parait bien assise sur le trepied du 
mendelisme, de la mutation et de la preadaptation, la verite oblige a ajouter 
des ombres au.tableau; je veux parler d’appareils dont le fini d’execution 
mis en regard de leur faible utilite apparente est difflcilement explicable 
dans l’une quelconque des theories de l’adaptation: ce sont les coaptations , 
ou ajustements reciproques de parties independantes, d’une curieuse per¬ 
fection mecanique, dont les Arthropodes, entre autres, nous offrent de; 
nombreux exemples. Natura maxime miranda in minimis. 


46 


L. CUENOT 


Avant (Taller plus loin, il est necessaire que je donne une breve descrip¬ 
tion de quelques-unes de ces coaptations. 1. Chez la plupart des Cephalo- 
podes Decapodes, le manteau est attache en deux points a la base de l’en- 
tonnoir, par lequel est expulsee Teau qui a passe dans la cavite palleale; sur 
Tentonnoir, se voient deux excavations dans lesquelles s’engagent deux saillies 
de forme parfaitement adequate, qui se trouvent a la face interne du manteau 
Cette attache elastique, tres solide, qui est du reste permanente pendant 
toute la vie du Mollusque, et ne peut etre disjointe que par une traction 
assez forte, correspond parfaitement au bouton-fermoir a ressort, ou bou- 




C 


Fig. 2. Belostoma: A, extremite distale de l’epimere mesothoracique droit vu par sa 
face dorsale, montrant le bouton saillant. 

B, bord de l’elytre droitvu par sa face inferieure, montrant la cavite. C, ou se loge le 
bouton. Gr. 10. 

ton-pression, imagine si je ne me trompe en 1886, dont on se sert pour fermer 
les gants et d’autres vetements. 

On retrouve le meme dispositif chez tous les Hemipteres aquatiques 
formant le groupe des Hydrocorises (Naucores, Nepes, Ranatres, Belostomes, 
Notonectes, Corises) pour attacher les hemelytres au thorax: Tepimere 
mesothoracique porte un bouton vaillant (fig. 2, A) qui s’engage dans un 
logement creuse a la face inferieure de l’hemelytre pres de son bord externe 
(fig- 2, B); quand on les separe Tun de Tautre en exer$ant une traction con- 
venable, on entend un petit claquement caracteristique, exactement comme 






GENETIQUE ET ADAPTATION 


47 


celui qu’on produit en defaisant un bouton de gant. L’adherence des 
deux parties est assuree avec plus de certitude par un pavage ecailleux 
qui revet les pieces chitineuses, exactement dans les regions coaptees, et 
qui fonctionne comme calage elastique. L’appareil est propre aux imagos 
et n’apparait qu’a la derniere mue, lorsque les hemelytres sont complete- 
ment developpes; sa Constance chez les formes aquatiques, et celles-la 
seulement, correspond sans aucun doute a une necessite de la vie dans 
l'eau; lorsque l’animal plonge, ses hemelytres restent etroitement colles au 
corps et ne se soulevent pas. 

2. Lorsque des pieces doivent glisser longitudinalement l’une sur l’autre 
d’un mouvement rapide sans se desunir, ainsi qu’il arrive pour des oviposi- 
teurs ou des aiguillons venimeux, elles presentent un assemblage tres 
ingenieux constitue par une languette saillante qui coulisse dans une rainure 
correspondante; la languette presente le profil d’un veritable rail a cham¬ 
pignon plus large au sommet qu’a la base, comme dans l’assemblage comme 
en menuiserie sous le nom de queue d’aronde. On en trouve de beaux 
exemples dans l’aiguillon des Fourmis, entre les deux stylets porteurs 
chacun d’une rainure laterale et le gorgeret porteur de deux rails, les parties 
coulissantes etant munies de petits ecailles chitineuses, fortement couchees 
vers la pointe de l’aiguillon, qui realisent un calage elastique. L’oviposi- 
teur des Locustes montre le meme assemblage, les valvules inferieures 
presentant chacune deux rainures, dans lesquelles glissent les rails des 
valvules superieures et internes. 

La meme coaptation, pour un tout autre but, se presente encore dans les 
elytres des Coleopteres; lorsque les elytres sont fermes sur le dos, il faut 
exercer une assez forte traction pour les separer; on les assemble de nouveau 
en les ramenant au contact sur la ligne mediane. Le bord interne ou 
sutural, toujours epaissi, de l’un des elytres, presente une languette saillante 
(fig. 3), qui regne sur toute la longueur, et s’engage a frottement assez 
dur tlans une gouttiere correspondante creusee dans le bord de l’autre 
elytre. A l’extremite anterieure des elytre, il y a presque toujours coapta¬ 
tion universe; le bord a languette presente un creuz dans lequel s’engage 
une saillie qui termine le bord a gouttiere. Mentionnons encore l’existence 
d’un calage elastique constitue par des asperites qui recouvrent les parties 
coaptantes (fig. 3, B). Grace a cet assemblage, les elytres fermes forment une 
cuirasse solide, difficile a disjoindre, qui recouvre exactement le dos du 
Coleoptere, et se coapte d’autre part avec les bords lateraux de l’abdomen. 
Il y a meme des cas ou l’adhesion est tellement forte qu’on ne peut plus la 
rompre, et que l’Insecte est dit a elytres soudes. La coaptation est nulle 
ou mal developpee chez les Coleopteres a elytres mous. 


48 


L. CUENOT 


3. Chez beaucoup de gros Lamellicornes, en particulier chez les Lucanes, 
les pattes anterieures presentent des coussinets de frottement (fig. 4), 
constitues par des brosses de poils: une de ces brosses se trouve sur la hanche, 
dans la region oii frotte le femur, et figure un triangle allonge; l’autre dessine 
un ovale sur la face frottante du femur. Ces coussinets manquent aux 
deux autres paires de pattes, et on ne voit pas trop quelle peut etre leur 
fonction; toujours est-il que grace a ces longs poils soyeux, le roulement du 
femur sur la hanche est d’une remarquable douceur. 




Fig. 3. A, schema de la coaptation des elytres de Coleopteres, a gauche le bord elytral 
porte une rainure ou coulisse la languette de l’elytre droit. 

B, coupe transverse du bord sutural des deux elytres, imago de Cassida viridis L. 
sortant de la pupe; p, pavage ecailleux assurant l’adhesion. Gr. 325. 

4. Les pattes revisseuses de modeles assez varies que Ton rencontre chez 
les Nepes, Ranatres, Naucores et Mantedes, presentent un caractere com- 
mun: a Tetat de repos, elles sont toujours fermees, a la maniere d’un 
couteau de poche, la partie correspondant a la lame (tibia + tarse) s’ap- 
pliquant d’une fagon exacte sur la partie correspondant au menche (femur), 
les courbures de l’une et de l’autre s’epousant parfaitement (fig. 5 et 7). 
Cet ajustement est tres curieux a examiner en detail: chez Nepa, le 
femur est creuse d’une forte gouttiere qu’on voit bien dans les coupes 






GENETIQUE ET ADAPTATION 


49 


(fig. 6); le tibia se pose sur les bords de la rainure, le tarse en forme de griffe 
ayant un petit logement special vers la base du femur; les bords des rainures 
femorale et tibiale portent une brosse de poils serres, plus grands et plus 




Fig. 4. Lucanus cervus L A, patte anterieure droite montrant les coussinets de 
frottement: c, coxa encastre dans le pro thorax, montrant Pextremite des poils du cous- 
sinet coxal;/, femur avec son coussinet ovale. Gr. 5. 

B , coxa vu de face, le femur etant enleve: s, surface triangulaire sur laquelle roule le 
femur, et dont la partie superieure porte le coussinet de poils; t, trochanter et articulation 
du femur. Gr. 6. 

forts que les autres poils du membre, et les deux pieces coaptantes ad¬ 
herent l’une a Pautre par Pintermediaire de ces poils qui s’entrepenetrent. 
C’est exactement ce qui se passe lorsqu’on appuie deux brosses Pune 
sur Pautre, poils contre poils; cet ingenieux dispositif permet aux deux seg- 







50 


L. CUENOT 



Fig. 5. Partie anterieure de Nepa cinerea L. montrant la coaptation des pattes ante- 
rieures ravisseuses; a gauche, la patte est dans la position de repos, le tibia rabattu sur le 
femur, et le tarse loge dans une encoche de la base du femur; a droite, le tibio-tarse est 
releve et laisse voir la rainure qui court sur la face anterieure du femur. Gr. 415. 




i 


Fig. 6. Coupe transverse de patte ravisseuse presque fermee, imago de Nepacinereas: 
b, brosses de poils sur les bords du femur (/) et du tibia (/). Gr. 40. 





GENETIQUE ET ADAPTATION 


51 


ments de rester appliques l’un sur l’autre sans qu’une effort musculaire soit 
necessaire. 

Chez Ranatra (fig. 7), la courbe concave du tibia est exactement adequate 
a la courbure convexe du femure sur lequel il se rabat; la rainure femorale 
qui occupe un peu moins de la moitie du segment estmathematiquementde 
la meme longueur que le tibia; a l’extremite proximale de la rainure se trouve 
une forte epine triangulaire derriere laquelle passe le tarse uniarticule, et 
qui forme pour ainsi dire un taquet d’arret. 



Fig. 7. Patte ravisseuse gauche de Ranatra linearis L.: c, coxa;/, femur portant une 
forte epine d’arret; t, tibia et tarse. Gr. 6. 

Chez Naucoris (fig. 8), la rainure femorale est bordee d’une galerie de 
courts piquants d’un cote, d’une brosse de poils de l’autre; le tarse d’un 
seul article qui continue la courbure tibiale est loge dans une depression 
faite expres, qui epouse exactement son contour. 

5. Les Coleopteres Longicornes presentent dans les deux sexes un ap- 
pareil musical, qui est constitue d’une part par une rape a stries transversales 
placee a la partie mediane du mesonotum, juste en avant du scutellum (fig. 
9), et d’autre part par une arete transverse situee a la partie interne du 
pronotum; par des mouvements de haut en bas du pro thorax, l’arete frotte 
contre la rape en produisant un bruit peu intense, en particulier lorsqu’on 
vient de capturer l’lnsecte. La rape, elegamment encadree d’une bordure 



52 


L. CUENOT 



Fig. 8. Patte anterieure ravisseuse de Nancoris maculatus Fabr.: /, femur volumineux 
et aplati; /, tibia; ta, tarse uniarticule; x, logement special de l’extremite du tarse. Gr. 40. 


? 



Fig. 9. Cerambyx miles Bon., vue dorsale de la rape musicale: e, bord sutural des 
elytres; m, mesothorax portant la rape entouree de son cadre; p , bord posterieur du pro¬ 
thorax, sous lequel se trouve l’arete de frottement; s, scutellum. Gr. 9. 













GENETIQUE ET ADAPTATION 


53 


lisse, a des nombreuses stries d’une extraordinaire finesse que Ton n’aperfoit 
qu’avec un eclairage special; Landois 7 en a compte 238 sur ]a rape du 
Cerambyx cerdo. On ne sait pas si cet appareil musical joue un role quel- 
conque dans la biologie des Longicornes. 

6. On pourrait citer encore maints exemples de coaptations, tous aussi 
remarquables que les precedents: Pappareil saltatoire ventral des Elaterides, 
la rainure qui se trouve le long du rostre des Curculionides et ou se loge si 
exactement Particle basilaire de l’antenne lorsque celle-ci se replie, les tres 
varies appareils d’accrochage des ailes pendant le vol chez les Rhynchotes, 
les Hymenopteres, les Lepidopteres, les crochets d’union des barbules dans 
les plumes locomotrices des Oiseaux, l’ajustement des femurs anterieurs 
des Phasmes et de la tete dans la pose de repos diurne (fig. 10), etc. 

II est prouve que les coaptations sont contenues en puissance dans le 
patrimoine hereditaire, car elles apparaissent deja chez l’embryon ou la 



Fig. 10. Vue de profil de la region anterieure de Carausius morosus, dans la position de 
repos diurne: a, base de l’antenne gauche s’engageant dans la gouttiere formee par les 
deux pattes anterieures; /, femur anterieur exactement applique contre la tete; p , pro¬ 
thorax; t, t£te. Gr. 3, 4. 

pupe, bien avant tout usage. Les deux parties qui, plus tard, seront coap- 
tantes, se developpent independamment Pune de Pautre et meme en des 
points eloignes, dans toute leur perfection; puis, au moment de l’eclosion, 
elles se deplissent, prennent leur forme, se rejoignent et s’ajustent exacte¬ 
ment du premier coup. Ainsi, pour ne prendre qu’un exemple, les pattes 
ravisseuses des Nepes et Ranatres sont chez l’embryon, non pas reployees 
dans la position qu’elles prendront chex l’animal eclos, mais completement 
allongees comme du reste les autres pattes; cependant on distingue tres bien, 
chez les embryons avances, la rainure longitudinale du femur et la petite 
encoche speciale ou viendra se loger le tarse. Des l’eclosion, lorsque la 
cuticle s’est deplissee et que le corps a pris sa forme definitive, bien qu’encore 
d’une fafon parfaite et commence a fonctionner. De meme pour les coap¬ 
tations speciales aux imagos, telles que le bouton-pression ou la rainure a 


7 Zeit. f. wiss. Zool., 17, 1867, p. 127. 











54 


L. CUENOT 


languette; les deux parties se forment chez la pupe, largement separees Tune 
de l’autre par la cuticule nymphale qui empeche tout contact; aussitot 
apres la mue imaginale, s’opere l’ajustage et la coaptation est apte a fonc- 
tionner des que la cuticule s’est durcie. 

Comment peut-on comprendre la genese des coaptations? Si insignifiants 
que paraissent ces minuscules details, cette question est une des enigmes de 
la Biologie. Ce qui est frappant, c’est que ces dispositifs ont souvent l’air, 
si je puis m’exprimer ainsi, d’avoir ete congus et executes par un ouvrier, 
c’est a dire que les solutions donnees par la Nature aux necessites des 
etres vivants sont des solutions analogues a celles que l’ouvrier humain a 
trouvees, par son intelligence propre, lorsqu’il a eu a resoudre des problemes 
a peu pres semblables (du reste sans copier la Nature, car la plupart du 
temps, il ignorait ce que celle-ci avait fait). La solution de la Nature ne 
differe de celle de l’ouvrier que par sa perfection bien plus grande, sa soup- 
lesse et sa solidite, son elegance et son luxe de petits details. 

Les solutions de l’Homme envisagent bien entendu un but ou un usage 
determine, ce que l’on appelle une fin intentionnelle en langage philoso- 
phique; les oeuvres de la Nature ont aussi une fin, qui est leur fonction, a 
laquelle elles sont pour la plupart parfaitement adaptees par leur structure, 
leur physiologie et leur coordination. Ainsi l’oeil a pour fin la vision. Que 
l’on dise que l’Homme voit parce qu’il a des yeux, au lieu de dire qu’il a 
des yeux pour voir, c’est un artifice de langage qui ne change rien au fond 
des choses. Cette finalite allait de soi dans la conception creationniste: 
Dieu, en creant chaque espece naturelle (la forme Sauterelle, la forme 
Hydrocorise), lui avait necessairement donne tout ce qui lui fallait pour 
vivre dans le milieu auquel elle etait destinee; le probleme des adaptations 
etait ainsi resolu d’un seul coup, en bloc. Mais l’avenement du trans- 
formisme a amene un changement complet dans Interpretation de la 
finalite: la grande majorite des biologistes, dans les cinquante ans qui 
ont suivi l’apparition du livre de Darwin sur YOrigine des especes , ont 
ete de purs mecanistes: si les organes ont une fin, disent-ils, c’est parce 
qu’ils remplissent un role utile, une fonction, dans l’organisme coordonne; 
ils ont atteint leur etat actuel par le jeu de causes naturelles (facteurs de 
revolution), la selection impitoyable n’ayant laisse subsister que des organes 
utiles ou d’une inutilite peu ou point genante, jamais d’organes nuisibles 
a l’espece. Les biologistes spiritualistes et les theologiens, tout en accep- 
tant revolution, croient a une causalite finale de l’Univers, a un Esprit 
createur superieur et anterieur a la Nature, qui dirige celle-ci par les lois 
qu’il a posees, mais sans intervenir directement; melior est causa causae 
quam causa causati , dit un vieil adage scolastique; ils conviennent que tout 


GENETIQUE ET ADAPTATION 


55 


ce qui se passe dans le monde releve de causes efficientes ou secondes, dont 
l’etude est l’objet de la science. 

Au point de vue philosophique, il y a sans doute un abime entre les deux 
concepts: pour les premiers, la finalite des organes est la resultante du 
hasard des variations et de la relection; pour les seconds, revolution est 
dirigee par la volonte d’un Esprit, suivant un plan preconqu et ordonne , 
par l’intermediaire des causes secondes; la finalite est done intentionnelle, 
qu’il s’agisse d’un microscopique rail de guidage d’un aiguillon d’Abeille 
ou d’un oeil d’Oiseau; pour le theologien, l’harmonie generate de la Nature, 
les merveilleux resultats des causes efficientes, constituent la preuve la 
plus apparente de la causalite finale. Mais au point de vue scientifique, 
les deux concepts se rejoignent absolument; pour expliquer la formation 
des coaptations, il importe peu qu’on soit spiritualiste, moniste ou agnosti- 
que, qu’on croie ou non a une causalite finale; on n’a le droit que d’en 
rechercher les causes efficientes (facteurs de revolution), exactement 
comme un physicien ou un chimiste qui etudie les phenomenes de sa 
speciality si on ne les trouve pas, il ne reste qu’a reconnaitre notre ignor¬ 
ance et a faire appel a un avenir mieux informe sur le nombre et la valeur 
de ces causes efficientes. 

De toute evidence, ce ne peut etre une variation germinale fortuite qui 
fait apparaitre d’un coup le bouton-pression et son logement, les deux 
brosses des pattes de Lucane, etc. On comprend bien qu’une mutation 
conditionne un changement de couleur, de symetrie, de nombre, de taille, 
de vigueur constitutionnelle, voire d’instinct, cest a dire quelque chose d’ 
indetermine au point de vue fonctionnel, sans relation avec l’utilite, dont 
l’animal s’arrange comme il peut; les innombrables mutations connues 
sont de ce type. Comment pourrait-on imaginer une mutation aveugle 
produisant d’un coup un resultat tel qu’une coaptation, aussi precise dans 
sa structure que dans sa fonction? 

Pense-t-on a une origine mecanique par pression et moulage reciproque, 
dont le resultat est devenu ensuite hereditaire? Outre que ce setait ad- 
mettre l’heredite d’un caractere mecanique acquis, ce qui est bien peu 
vraisemblable, cela n’expliquerait rien; beaucoup de coaptations sont des 
structures beaucoup trop raffinees pour etre comparees a des accidents 
d’origine surement mecanique, comme les sillons grossiers que creusent a 
la surface d’un os la pression d’un vaisseau ou le frottement d’un tendon. 
Comment la rape mesothoracique d’un Longicorne et son frottoir pro- 
thoracique, les appareils d’assemblage des ailes et des elytres, le rail et sa 
rainure, pourraient-ils etre le resultats de pressions, de frottements ou de 
tout autre acte mecanique? Ce serait incomprehensible, d’ autant plus que 


56 


L. CUENOT 


beaucoup de ces coaptations, dans le cas special des Insectes, sont propres 
aux imagos, dont le tegument est fixe une fois pour toute et ne peut plus 
6tre deforme; il en est aussi qui ne remplissent un role que pendant quelques 
minutes de la vie (ovipositeur des Sauterelles). Admettre qu’une coapta¬ 
tion, ou un organe comme l’oeil, ou une forme comme celle du Poisson avec 
ses courbures, ses plans stabilisateurs et ses nageoires, provienne des jeu de 
forces naturelles agissant sur la matiere organique, sans agent directeur, c’est 
attribuer a celle-ci une capacite sui generis de repondre a des excitations 
simples par l’edification mysterieuse de machines complexes et coordonnees, 
ce qui nous remene au finalisme d’Aristote ou peu s’en faut. 

II ne reste qu’une hypo these je ne dis pas satisfaisante mais possible: 
ies coaptations seraient le resultat d’une evolution dirigee par l’utilite, a 
partir d’organes tres simples, mais deja utiles; toutes les variations qui se 
transmettront realiseront forcement des ameliorations de l’etat primordial 
ou au pis aller des modifications indifierentes, car toutes les variations de- 
favorables determinent la mort des individus qui les presentent et s’etei- 
gnent. II est evident qu’un simple defaut a une glissiere d’ovipositeur em- 
p6che absolument la ponte, d’ou elimination certaine de la lignee desavan- 
tagee; il est possible que la coaptation des pattes ravisseuses, voire meme 
que les coussinets de poils des pattes de Lucanes ou la rape musicale des 
Longicornes, aient une importance que nous ne soup^onnons pas; nous 
sommes tres mal places pour juger de l’utilite de ces petits details pour la 
vie de l’espece. La morphologie comparee des coaptations dans un groupe 
naturel n’est pas defavorable a cette maniere de voir; il y a chez les Locustes 
a ovipositeurs courts des glissieres assez frustes mais qui suffisent dans 
le cas particulier; dans le groupe des Hemipteres aquatiques, il y a des 
appareils beaucoup moins parfaits que le bouton-pression (Corises), mais 
qui fonctionnent neanmoins. L’oeil humain, qui est assurement un eton- 
nant appareil d’optique ayant la vision comme fin, derive d’une simple tache 
pigmentaire innervee; dans la lignee des ancestres vivant a la lumiere et 
dependant d’elle pour chercher leur nourriture a la vue ou pour se proteger, 
toutes les variations qui ont pu se produire autour de l’oeil ou a son inter- 
ieur n’ont realise que des etapes graduees a la fois dans la structure et dans 
1’utilite, ou au pis que des modifications indifierentes (comme les variations 
de couleur de l’iris); toutes les variations defavorables, produisant une 
diminution de la vision, ont abouti a la decadence et a la mort des especes, 
et nous ne nous les connaissons pas, ou bien les ont contraintes a chercher 
un milieu obscur, ou elles pouvaient se passer de la fonction (Poissons aveu- 
gles, Reptiles en doges, Taupe, etc.). 


GENETIQUE ET ADAPTATION 


57 


Sans doute c’est attribuer une grande valeur a la selection, que Ton a 
une tendance a notre epoque a sous-estimer, et a reduire a un role conserva- 
teur. A cela on peut repondre que son importance est prouvee avec une 
rigueur presque experimentale par la cessation meme de son action, ce que 
Weismann a appele la panmixie; ce processus est bien connu d’une part 
chez les animaux domestiques proteges par l’Homme et d’autre part chez 
l’Homme lui-meme, soustrait par sa vie sociale a la lutte contre la faim, le 
froid, les carnassiers, et aussi par la medecine et les institutions charitables 
aux consequences des malformations et des maladies microbiennes. Les 
premiers renferment des quantites de races (Chiens de luxe, animaux gros, 
Oiseaux incapables de voler) qui ne vivraient pas une semaine a l’etat de 
nature; le second, surtout a l’etat civilise ne compte plus les caracteres 
presque pathologiques des dents, des poils, des glandes mammaires, des 
yeux, etc., qui auraient rendu sa vie impossible dans les ages prehistoriques. 
II n’est done pas invraisemblable d’admettre que la selection a une in¬ 
fluence assez grande pour diriger F evolution de petits appareils qui ne sont 
peut-etre insignifiants qu’en apparance, et qu’elle puisse jouer le role de 
l’ouvrier qui perfectionne graduellement son oeuvre. 

J’ai expose loyalement, je pense, la these des selectionnistes; je dois 
dire que, pour ma part, elle ne me satisfait pas pleinement. J’ai peine a 
croire que des mutations de hasard du patrimoine hereditaire, meme canali- 
sees par la selection, puissent edifier un organe complexe impliquant un 
nombre enorme de coordinations, tels que l’oeil et certaines coaptations. 
Je trouve incomprehensible que des etres aussi differents qu’un Pecten, 
qu’un Cephalopode et qu’un Poisson, aient acquis des yeux d’une perfection 
comparable, par le simple jeu de variations fortuites superposees (argument 
de Bergson, dans VEvolution creatrice, 1907); c’est d’autant plus incroyable 
que si les yeux sont indispensables au Cephalopode et au Poisson, qui 
chassent a la vue, le Pecten qui se nourrit passivement pourrait fort bien 
s’en passer, comme tant d’autres Lamellibranches. 

Quant a l’importance attribute plus haut a la selection, c’est pure hypo- 
these il est vrai, mais on ne saurait positivement la critiquer, car on est bien 
peu fixe sur la valeur du facteur selectif. D’un cote l’on voit des variations 
considerables (especes polymorphes) qui subsistent cote a cote sans qu’il y 
ait entre elles de concurrence visible; d’un autre cote, on connait le cas de 
YAmphidasis betularia dont la variete melanique carbonaria est en voie, 
depuis 70 ans, de se substituer au type tachete dans maintes regions; ailleurs 
encore la selection a un role conservateur, en eliminant les individus les 
plus eloignes de la moyenne raciale. Nous ne savons pas du tout si, toutes 
choses etant egales, une legere mutation favorable d’un appareil coaptatif 


58 


L. CUENOT 


peut constituer un avantage qui donne prise a la selection, ou bien si cela se 
perd dans la masse des contingences. 

Peut-etre a cause de ces incertitudes, on a le sentiment obscur qu’il manque 
quelque chose a nos conceptions de revolution et a ses causes efficientes; 
a tort ou a raison, les mutations fortuites et la selection, qui sont les seuls 
agents reconnus de perfectionnement, nous paraissent insufiisants. La 
necessity d’un facteur nouveau, interne ou externe aux organismes, regula- 
teur des mutations et capable de les diriger vers une fin, se fait surtout sentir 
lorsqu’on etudie des organes complexes comme les yeux, les organes photo¬ 
genes des animaux abyssaux, les organes electriques des Poissons ou les 
coaptations, ou lorsqu’on contemple les ornementations d’une si rare et 
d’une si inutile beaute des plumes du Paon et de V Argus. Du reste, je n’en- 
tends par la rien de metaphysique, aucune entelechie aristotelienne ou 
leibnizienne, mais quelque loi generate, qui a echappe a nos predecesseurs, 
et qui reste a decouvrir. 


INHERITANCE IN UNICELLULAR ORGANISMS 

H. S. JENNINGS 
The Johns Hopkins University 

Of the results of the study of inheritance in unicellular organisms, I 
can attempt only a summary of certain matters that appear of general 
interest, particularly to those occupied primarily with the problems of race 
improvement. 

In man and in all higher animals the problem of racial alteration, and 
indeed all problems of genetics, are enormously complicated by the mixture 
of two individuals at reproduction, accompanied by a sorting over and 
recombination of the materials on which the diversity of characteristics 
depend. This results in a tremendous diversity among the descendants, 
purely as a matter of kaleidoscopic regrouping; it becomes extremely difficult 
to distinguish among these any actual racial changes, any changes due to 
alterations in the hereditary substances; difficult to study any of the 
phenomena of inheritance except those resulting from this sorting over and 
regrouping. In consequence, inheritance under the name of Mendelism has 
largely come to be identified as simply the results of this sorting over and 
recombination. 

In unicellular organisms this sorting and recombining is omitted for 
long periods; descent for thousands of generations is from a single parent; 
all the progeny have the. same germinal material or “genes” as had the 
parent. We are therefore enabled to observe what inheritance is when 
quite free from Mendelism; we may examine any hereditary changes due 
to alterations in the germ plasm, quite uncomplicated by kaleidoscopic 
regrouping of the germinal substances. 

When we follow for long successions of generations the descent of organ¬ 
isms multiplying thus from a single parent, two impressions are very 
strong. First, we are astonished at the uniformity, the constancy, shown 
by the hereditary constitution of the race. One is almost led to believe 
that it is a fixed, an unchangeable, thing; some investigators that have 
devoted themselves for years to this work still hold that it is an unchanging 
thing. Second, in spite of this uniformity of the hereditary constitution, 
we are struck by the variety, the sometimes very great diversity, among the 
individuals all possessing this same hereditary constitution; all arising 

59 


60 


H. S. JENNINGS 


from the division of a single parent. To take the case first worked out, in 
Paramecium there are many races differing very slightly in size, and these 
differences are inherited completely, if we compare the mean sizes of the 
different races. In each race taken by itself there are similar or greater 
differences in size among the individuals. But these differences are not 
inherited at all; large and small individuals of the stock produce progeny 
not differing in the least. In some other protozoa the characteristics are 
more diversified. In Difflugia there are similar diverse races, differing in 
mahy structural features, and within the single race we find the same 
structural diversities; differences in the number and arrangement of the 
spines; in the length of the spines, and the like. But these differences 
within the race appear not inherited; an individual with many spines and an 
individual with none produce progeny of the same average character, if 
they belong to the same stock; and so of other characteristics. We have in 
this respect a parallel to the spotting, to the distribution of colors, in some 
mammals; the individuals may differ in most characteristic ways, but the 
differences are not inherited. Yet, in our Difflugias, and in Paramecium, 
identically the same differences, when characteristic of different races , are 
inherited. 

This great diversity of individuals all possessing the same hereditary 
constitution appears worthy of particular notice by those interested in the 
improvement of a species. There is a popular impression that genetics, 
that eugenics, is attempting to teach that our characteristics—that all 
that we are—are predetermined in the germ plasm; that with the germinal 
constitution which we receive all our characteristics—our fate—are decided. 
This is not true in the Protozoa; I venture to say that there is no ground 
for asserting it to be true in man. Merely because we inherit a certain 
characteristic, we are not required to have it. In our Protozoa, individuals 
with the same hereditary constitution possess most diverse characteristics. 
It is not inherently impossible that from the same stock of Difflugia, all 
with the same hereditary constitution, we could produce two sets—one 
exhibiting always numerous large handsome spines, and producing progeny 
with such; the other always with few, stunted, or lacking spines, for these 
diversities are seemingly not inherited,—though we do not know just how to 
do it yet. 

But not all characteristics show this “personal” diversity, independent 
of the germinal constitution. In Paramecium, the form and structure do 
not show it; they seem always fully determined by the hereditary constitu¬ 
tion; while size does show it. In Difflugia, the number, size, and arrange¬ 
ment of spines is thus largely subject to personal variation, while bodily 


INHERITANCE IN UNICELLULAR ORGANISMS 


61 


size is not; body form is not; size, structure, and number of teeth are not. 
In every organism it is necessary to determine by observation which 
characteristics are fixed by the germinal constitution, which ones are not; 
and there are very great diversities in the degree of fixation by inherited 
constitution, in the readiness to be modified by the environment, in different 
characteristics of the same race. 

Though the individuals of a race thus vary, there is, as we remarked, a 
high degree of constancy, of uniformity, in a given germinal constitution; 
experiment for thousands of generations may reveal no alteration in it. 
There are many lower organisms that have thus been thoroughly studied 
for long periods while reproducing from a single parent, in which no single 
diversity in hereditary constitution has been discovered. There are 
indications, as we know, that in man and higher animals the same sort of 
reproduction would show similar constancy and uniformity. When two 
individuals are derived from a single egg we have essentially the same 
phenomenon as in production from a single parent; they receive the same 
germinal material and are now in physical features almost exactly alike, 
forming the so-called identical twins. In the armidillo, as Newman and 
Patterson show, several offspring arise from a single egg and these show the 
same astonishing similarity that we see in identical twins in man. In the 
Protozoa, identical twins occur in sets of millions each. In man, as identical 
twins show, form and structure appear to be very accurately fixed by the 
germinal constitution rather than by the action of the environment, while 
behavior and physiological processes are not. 

Yet permanence of inherited constitution turns out not to be the whole 
story after all, in reproduction from a single parent. The “personal” 
differences among members of the same stock, apparently quite non- 
heritable, turn out, on prolonged study to be not absolutely so. Breeding 
organisms particularly favorable for such study, where growth changes 
and environmental alterations during life are excluded, and following 
uniparental reproduction for long periods, we find in some cases that after 
a great number of generations slight hereditary differences have appeared. 
Selecting on the one hand individuals possessing a certain character, on 
the other those not possessing it, we find that after a long period we have 
obtained, from a single parent, stocks differing hereditarily in these respects; 
the differences persist from generation to generation. So in the rhizopod 
Difflugia I finally obtained as descendants of a single parent a number of 
stocks showing different combinations of diverse hereditary characters. 
Some were large with large spines; some large with small spines; some small 
with large spines, and the like. Similar inherited differences have been 


62 


H. S. JENNINGS 


shown to arise in Arcella by Hegner, and in Centropyxis by Root; inherited 
physiological diversities have been shown by Middleton to arise in infusoria. 

It appears thus that the “personal” differences within a stock are not 
after all entirely non-heritable, as they appear to be if we study them for a 
few generations. There is in the very long run a low degree of correlation 
between the “personal” characteristics of the parent and those of the 
offspring, even within the same stock. Here we have a phenomenon of 
inheritance quite apart from Mendelism. So far as it goes it tends to 
rehabilitate the Galtonian theory of inheritance as a correlation between 
the bodily characteristics of parent and offspring, quite independently of 
chromosomal recombinations. 

It must not however be passed over in silence that in many organisms, 
multiplying by uniparental reproduction, after long study, no inherited 
differences are found to occur in a stock; the hereditary constitution appears 
permanent. 

Where inherited diversities do occur, where do they arise, and how are 
they brought about? 

On this we know in the higher Protozoa very little. In Arcella, Hegner 
found that the number of nuclei may be increased by a division of a nucleus 
without a division of the body, or decreased in the reverse way. Such a 
change in the number of the nuclei induces a change in the size of the cell 
body, and correlated with this other changes occur; the number and size 
of the spines change. Thus differentiated races arise sometimes through 
these changes in the number of the nuclei. It is clear that not all the 
inherited variations arise in this manner, for there are in Difflugia diverse 
combinations of characters; large races with large spines; other larger 
races with smaller spines. Aside from this, there is little but speculation 
as to the origin of the hereditary differentiations. 

May diversities in the environment bring about hereditary differentia¬ 
tions? Here again the conditions are parallel to those already described. 
In many cases environmental agents have been tested for many generations; 
they readily modify the individuals on which they act; but as soon as the 
original environment is restored, all these modifications are lost; there is 
no “ inheritance of acquired characters.” Such has been my own experience 
in extensive experiments with chemicals on Paramecium. Such is the 
large, obvious fact in most work of this sort. Other investigators have 
been more successful. Middleton found that when a part of the progeny 
of a single individual of Stylonychia are kept at a high temperature, the 
others at low temperature, the two become permanently diverse in rate of 
fission. The fact that we are here apparently dealing with inherited 


INHERITANCE IN UNICELLULAR ORGANISMS 


63 


differences in what may be called vitality makes doubtful the bearing of 
these observations on characteristics of other sorts. Middleton has recently 
observed similar permanent changes as a result of subjection to chemicals. 
By subjection to chemicals for long periods, Jollos has induced hereditary 
changes of resistance to these chemicals in Paramecium. The greater 
resistance induced lasted for only a limited number of generations as a rule, 
but the longer the given chemical acted the longer did the inheritance last. 
This would suggest that if the action of the environmental agent were very 
long continued, the inherited alteration might become as permanent as 
any hereditary character, though Jollos doubts that this would occur. 

If we turn from what we may call the higher , and the free-living, uni¬ 
cellular organisms to the simpler ones, the picture appears to change 
greatly, with relation to the inherited effects of environmental action. 
In the bacteria and some of the pathogenic protozoa, great numbers of 
cases of inherited modifications due to the environment are reported; 
changes in resistance, in virulence, and the like. Here the difficulty of 
being certain that we are dealing with progeny of a single individual is very 
great, so that many hold that these results are largely deceptive. It is 
asserted that the experiments began with a mixture of races of diverse 
hereditary characteristics. Some of these thrive under certain conditions 
and therefore give character to the cultures; others thrive under other 
conditions, giving a new character to the cultures; this alone is held to be 
the explanation of the apparent change in hereditary characteristics due to 
environmental differences. There are investigators who urge strongly 
this interpretation for all cases. 

As this is a field in which my own work has not lain, I shall not take it up 
in detail but will make the following general remarks. First, in some of the 
investigations the evidence appears very strong that the observed hereditary 
changes are actually the result of environmental modifications. Second, 
if we accept generally the accounts of the phenomena in bacteria, as set 
forth for example in Adami’s “Medical Contributions to the Theory of 
Evolution, ” we must conclude that in these elementary organisms hereditary 
modification due to environmental action is rather readily produced; and 
that it is adaptive in character; that here in the lowest organisms “acquired 
characters are inherited ” on a rather large scale, and must be an effective 
factor in evolutionary change. These conclusions I think have a very 
great body of evidence for them. If they are correct, the ground for the 
difference in these respects between them and the higher unicellular organ¬ 
isms, as well as multicellular organisms, becomes a problem of great interest; 
one is inclined to believe that the difference must be one of degree only. 


64 


H. S. JENNINGS 


Biparental reproduction occurs of course in many of the unicellular forms 
but its results have been little studied. There is some evidence that it 
produces diversity of the same sort seen in higher animals, through the 
sorting and recombination of the germinal materials. 

In sum, the study of inheritance in unicellular organisms permits us to 
obtain and examine great numbers of individuals having identically the 
same hereditary constitution. “Personal” differences of marked character 
occur among such individuals, emphasizing the fact that heredity by no 
means fully determines the characteristics of organisms; all characters 
are due to the interaction of germinal materials with the other conditions, 
and are dependent on both. A remarkable constancy and uniformity of 
the hereditary constitution itself is discovered; a constancy so great as to 
give the impression that no heritable changes occur. But in some of 
these organisms, in the long run, it is found that the diverse “personal” 
features have a faint tendency to be inherited; gradual changes in hereditary 
constitution occur in the passage of a great number of generations. In 
the lower unicellular forms, particularly the bacteria, such hereditary 
change appears to result from the action of the environment, “acquired 
characters” being inherited, and that not so very uncommonly. But this 
conclusion is rendered somewhat insecure by the difficulty of determining 
critically the facts in these minute organisms. In the more complex 
unicellular forms, as in the multicellular forms, hereditary change resulting 
from environmental action appears rare if not absent. 


EVOLUTION OF THE CHROMOSOME COMPLEX 

C. E. McCLUNG 
University of Pennsylvania 

It is significant of the development of modern biology that a cytologist 
should be invited to discuss the nature of chromosome organization and 
descent at an eugenics congress. By this circumstance there is shown a 
real recognition of the relation existing between the material substratum 
of hereditary processes and their visible somatic manifestations. This 
approach between fields which, two decades ago, had little in common is 
most gratifying and its fruitfulness is shown in the many important genetical 
advances made each year. Correspondingly it becomes increasingly 
important to determine accurately the character and range of the cytological 
phenomena by which genetical results are thus explained. 

The chromosome theory of heredity is now so firmly established that only 
an entire revolution in our conception of cellular organization can displace 
it as a formal explanation. As new facts accumulate in the study of in¬ 
heritance they fall at once into relation with others because there now 
exists an adequate conception of the germ cell mechanism. Facts, at 
first thought to discredit the chromosome theory, have, with increased 
knowledge, been found to relate to special conditions whose explanation in 
turn has broadened the original conception. Thus, for biology, the chromo¬ 
some theory serves to correlate and promote advances in knowledge much 
as the atomic theory did for chemistry. 

Since the significance of an evolution of the chromosome complex is 
bound up with the functions of the chromosomes in individual development, 
it is important to recall our understanding of the chromosomes in this role. 
It is again a question of the relation between ontogeny and phylogeny. 
Briefly stated my conception is as follows: The chromatin is that portion 
of the cell most immediately concerned with the control of processes which 
result in the formation of structures and the exhibition of functions which 
are called characters. This chromatin is differentiated into structural 
units, each of which has a particular function to perform in relation to the 
series of functions inherent in the entire chromatin of the cell. These 
units are grouped into chromosomes, in which, at their greatest extension, 
they are linearly arranged. The chromosomes in number, form and size 

65 


66 


C. E. MCCLUNG 


are constant for the species and thus give evidence of the specific character 
of their constituent parts. Individually they show constant differences 
within the cell which is taken to be an indication of the unlikeness of their 
constituent units. 

In the present state of our knowledge it is hazardous to attempt the 
formulation of theories of chromosome relations between widely different 
forms of animals. It is much safer to enquire into the conditions obtaining 
within limited and well-understood groups. With this thought in mind I 
purpose to review briefly those facts which have developed during the last 
two decades in extensive studies made by myself and students upon the 
chromosomes of certain Orthopteran groups. With these will be compared 
the results of other investigations which seem to contribute to a wider 
knowledge. 

The outstanding fact which has been revealed by this study of practically 
all of the genera of North American Acrididae is that the entire family 
possesses a characteristic chromosome complex, the most marked feature 
of which, in nearly every case, is a common number. It is difficult to 
over-estimate the significance of such a fact in relation to the phylogeny of 
the group. There are scores of genera, hundreds of species, billions of 
individuals, all containing multitudes of cells, each of which possesses the 
same numerical complex of chromosomes as every other cell in all the in¬ 
numerable host. And what is true of the living population we must assume 
to have prevailed in the endless generation of past geological ages, because 
this uniformity in members of divergent terminal series could not eventuate 
from disharmonious ancestral groups. It passes the imagination to con¬ 
ceive an organization of material so exact as to accomplish this endless 
uniformity and yet, in every mitotic division, is exhibited the means by 
which it is done. Each chromosome there visibly reproduces itself. The 
chromosomes now existing are the direct lineal descendants of similar ones 
down through the ages. There is absolutely no logical escape from this 
conclusion in the light of existing evidence. However much conditions 
may vary in the other groups, within the Acrididae we have this 
demonstration of the phylogenetic continuity of the chromosome complex. 
But if the chromatin is indeed the differentiated congeries of character 
determiners whose nature is, in part, revealed by the form of integration 
into chromosomes, then we should find, in various species, definitely recog¬ 
nizable chromosome distinctions. These, indeed, occur and are of the most 
exact taxonomic significance as will later appear, but, so far as the totality of 
the complex is concerned, practically all the evidence speaks for it. In 
species after species the male animal shows, in every cell, a group of twenty- 
three chromosomes while the female cells have always twenty-four. 


EVOLUTION OF THE CHROMOSOME COMPLEX 


67 


Not only are there always the same numerical series of chromosomes in 
these Orthopteran cells, but they occur in an approximately like size 
seriation, ranging through proportionately equal steps from the smallest 
to one five to ten times its size. Despite a considerable range in actual 
dimensions of chromosomes in different species, there is always found this 
gradation from the smallest to the largest. How far this criterion may be 
used to determine actual homologues throughout the taxonomic group is 
as yet undetermined, but the weight of evidence would indicate its value 
in a general way. If it were exact we could, of course, homologise any 
well defined element like the smallest chromosomes in all the species of the 
family. Whenever such definite identification becomes possible our means 
of structural analyses will be greatly increased. As yet the only definitely 
recognizable chromosome in all the species is the accessory chromosome. 
This may be identified beyond any question in every one of the many 
individuals studied. 

It is clear from these considerations, therefore, that, although we are 
dealing with the same series of chromosomes they must be individually of a 
different internal constitution in different species if they are indeed the 
groups of determiners we conceive them to be. Such internal differences 
may or may not be visible in the general configuration of the element, and, 
as yet, we have no means of measuring such differences so that they may be 
correlated with taxonomic characters. Although these internal changes 
may not with certainty be determined there are species whose chromosomes 
show by their behavior that they have undergone some structural modifica¬ 
tion. Perhaps the most marked instances of this are where an actual 
difference in the number of chromosomes appears to exist. In the Acrididae 
of North America, which we have so far studied, this numerical variation is 
principally due to fusions between certain non-homologous members of the 
complex, which also breaks the size seriation. Studied alone and without 
consideration of other members of the group, such cases as have been de¬ 
scribed for Hesperotettix, Mermiria, Chloealtis, and Chorthippus would 
seem to be evidence against the generalization of a family constancy for 
the chromosome complex, but, when properly understood, are found to be 
merely instances of a chromosome reorganization which has brought 
about a more or less temporary union between certain of its members. 
Since this tendency to form multiples resides definitely in certain taxonomic 
groups we must regard this behavior of the chromosomes involved as an 
evidence of a peculiarity in their organization. That it is perhaps of the 
same nature wherever found is suggested by the fact that the euchromosome 
multiples tend to involve the largest members of the complex and that 
additional ones follow down the size series. 


68 


C. E. MCCLUNG 


While in the Acrididae evidences of internal reorganization manifested 
by numerical and size changes are generally confined to a reduction in 
number by fusions, in other groups it is found that the total series of 
chromatin elements may be maintained in the presence of marked numerical 
chromosome variations. Thus it has been found by Holt that the type 
number of six in Culex may be multiplied in a single cell to seventy-two 
by successive duplications of the individual chromosomes. The diploid 
number of forty in the pig may be increased to fifty-eight by fragmenta¬ 
tion, according to Hance. By non-disjunction of homologues it is found by 
Bridges that the number in Drosophila may be raised or lowered by one. 
In various forms there are also multiple complexes, and supernumeraries. 
Blakeslee and his fellow workers have noted twelve distinct mutants of 
Datura, each of which shows a different triple representation of homologous 
chromosomes in the diploid complex making 25 instead of 24. A thirteenth 
mutant has four homologues or a tetraploid complex, and it differs from all 
the others. 

Wherever these variations occur, however, it is found that the total 
normal series of chromatin elements is not reduced, although there may be 
combinations, duplications, additions or rearrangements. The series, in 
whole or in part, tends to maintain itself. This is especially true of the 
germ cells where it would appear from all evidence, genetical and cytological, 
there must be maintained a full complement of the chromosomes. Addi¬ 
tions may be made to this normal number of all members of the complex, 
but it may not, without injury, be reduced. Distinct somatic differences 
sometimes accompany such modifications of the germ cell chromosomes, but 
the changes are not destructive of the coordinated action of the complex. 
On the other hand there is some evidence to indicate that somatic cells 
may suffer more extensive chromosome modifications without injury to their 
specific action. 

While the conditions of the chromosomes in the Acrididae are thus 
comparable throughout the family, on the basis of our present knowledge, 
comparisons between even the saltatorial families of Orthoptera are not at 
present possible. It is here that we may hope, with increased knowledge, 
to gain a conception of the phylogenetic history of a chromosome group. 
Already the accessory chromosomes may be homologised definitely wherever 
found. If it is true that in one element of the complex we have a definitely 
integrated portion of the idioplasm, the presumption is strong that there 
may be other recognizable chromatin units. That the Acrididaean numbers 
23 and 24 appear in the Gryllidae may not be without significance, therefore. 

How the variations in number of chromosomes, marking the difference 
in the form of chromatin organization came about, we have no direct 


EVOLUTION OF THE CHROMOSOME COMPLEX 


69 


indication. It may be noted, however, that the changes in the Acrididae so 
far observed are all concerned with reducing the number of free units by 
combinations to produce multiples. Sometimes these reductions are 
permanent as in Chloealtis, Chorthippus, etc.; sometimes they are occa¬ 
sional, as in Hesperotettix viridis. That such combinations are indicative 
of a definite tendency is shown by their prevalence in numerous species of 
the genera concerned. Also it is significant that such combinations begin 
with the largest pair and successively involve the series in this order, so 
long as multiples are formed. These facts indicate that some general 
principle of chromosome organization is expressed in this tendency of 
chromosomes to form unions. When this does occur, however, there is 
a break in the uniform series of chromosome sizes commonly found in a cell, 
and we should expect to discover some later reorganization of the elements 
in case of permanent fusions. This does not seem to be manifest in the 
species so far studied. One other change in chromosome structure may be 
important. In most species of Acrididae, the attachment of fibers to the 
chromosomes in the metaphase is terminal but occasionally, as in Circo- 
tettix and Trimerotropis, it may be median or subterminal. Since fiber 
attachment is so constant a feature of chromosome structure this change is 
also suggestive, particularly in view of the variable taxonomic character of 
the groups of species involved. 

The conclusion that these rearrangements in the relations of the chromatic 
substance found in the Acrididae may be indicative of a principle con¬ 
cerned generally with evolution or change in the chromosome complex is 
strengthened by observations that similar conditions obtain in the Locusti- 
dae. In the genus Jamaicana, Miss Woolsey finds the formation of multiples 
similar to those of Hesperotettix and Chorthippus, and earlier I had noted 
a multiple involving the accessory chromosomes in Anabus, similar to those 
in Hesperotettix and Mermiria. The relatively frequent involvement of 
the accessory chromosome in multiples may be an indication of change in 
its structure, because it is clear that it carries other determiners than those 
involved with sex differences. This element, because of its alternate 
passage from male to female, its differential rate of metabolism and union 
with certain euchromosomes is indeed admirably fitted to act as a means of 
promoting variations within the material basis of heredity. 

In attempting a correlation between chromosomal conditions and the 
exhibition of body characters, it must be borne in mind that the differences, 
for instance, between the various saltatorial families of the Orthoptera, are 
not large, relatively. They pertain much more to relative degrees in 
development of a common series of parts than to the presence or absence of 


70 


C. E. MCCLUNG 


unique structures. Our problem is then, not to find such additions or 
subtractions of chromatin substance as would account for the appearance 
or disappearance of distinctive bodily structures, but rather to note such 
changes in the hereditary mechanism as would explain proportionate 
increases or decreases in development of a like series of structural elements 
or characters. Part by part grasshoppers, locusts, or crickets are not so 
much unlike except in details of form, size, or proportion. 

Indeed, considering the characteristics of the Acrididae as compared 
with those of the insect type in general the differences are not great. Func¬ 
tion by function the activities of insects are performed surprisingly alike 
throughout the various Orders. Organs of perception of movement and of 
metabolism differ only in details of structure, not in fundamental character. 
It would be unreasonable to expect in the individual, therefore, any pro¬ 
found differences in the nature of the material substance controlling the 
development of the characters marking the group. Provided the ultimate 
units or genes were present in the germ cell, in their entirety, they might 
experience considerable variation in their form of integration without 
producing very marked somatic distinctions. It is not surprising, therefore, 
to find that rather marked differences in form, size, and behavior of chromo¬ 
somes are only of generic significance. 

Instances of distinct generic differences are not uncommon in the Acri¬ 
didae and they concern many features of chromosome organization. Thus, 
in Mecostethus, all the chromosomes are of a slender rod shape and in the 
first spermatocyte lie extended in the equatorial plate without the formation 
of rings. Such a complex has not been found elsewhere among all the 
species studied. Chloealtis exhibits a first spermatocyte complex in which 
the largest three pairs are united into multiples in the form of plain rings 
lying parallel to the spindle axis. Chorthippus shows a similar arrange¬ 
ment except that the rings are extended in part also within the plane of 
the equatorial plate. Hesperotettix viridis presents in different individuals 
a considerable variation in the union of chromosomes into multiples, and 
some of these conditions appear in other species of the genus. In Circo- 
tettix is found so permanent a union between two chromosomes that the 
normal number is apparently reduced by one. Accompanying this numer¬ 
ical difference there are shifts in the point of fiber attachment in many of the 
chromosomes so that they appear much different in shape. 

Variations of this character are relatively of considerable magnitude and 
must indicate a measurable change in the nature of the ultimate chromatin 
units, and yet their effects, judged by taxonomic characters, are only of 
generic value. So far they are of undefinable significance in determining 
the general nature of chromatin changes that occur when organisms vary. 


EVOLUTION OF THE CHROMOSOME COMPLEX 


71 


If, therefore, we lack understanding of the meaning of the strongly 
marked chromatin differences in a group of organisms so well worked 
relatively as the Acrididae, it would be of little profit to attempt appraise¬ 
ment of much less definite and marked variations in the chromosomes of 
widely removed groups of organisms. More than a thousand species of 
animals have now been studied, and it would seem that if there are signifi¬ 
cant evolutionary differences visible in the chromosomes they would have 
been detected. It was suggested by Meek that the diameters of the chromo¬ 
somes increase in a definite way according to the evolutionary development 
of animal groups, but this was so patently incorrect that he was himself 
obliged to repudiate the suggestion. Attempts have been made to show 
that large chromosome numbers are characteristic of low development or 
specialization, but the criterion does not hold at all, for both small and large 
chromosome numbers are found in various groups of the most simple and 
of the most complex organization. Undoubtedly these differences in 
number are significant because they indicate variations in the grouping of 
the ultimate chromatin units, but, we lack as yet criteria for determining 
their meaning. 

From these general studies upon chromosome numbers there have, how¬ 
ever, appeared some facts of importance. They indicate, for instance, 
that even in groups with wide numerical ranges, there is a chromosome 
number of most frequent occurrence, and, in many cases, other numbers are 
multiples of this. Mrs. Harvey has called attention to the fact also that 
in groups showing marked affinities like the molluscs and annelids, the 
type numbers are the same. Although actual numbers may vary from 
one to a hundred, the range of these type numbers for Phyla is not great. 
Commonly the number does not exceed eighteen and generally falls below 
twelve. Every such evidence of uniformity in the integration of the 
chromatin speaks for unity in the form of chromosome organization and 
thus for continuity through phylogenetic history. 

Important evidence of the relations existing between the chromosomes 
of different species within a group, where there is a range of chromosome 
numbers, is afforded by the studies of Metz upon species of Drosophila. 
Judging by relative size and the point of fiber attachment, he is able to 
show how two rod shaped chromosomes of one species are represented by 
a single V-shaped element in another. 

If the complex of chromosomes were absolutely constant throughout a 
group of organisms, the determination of homologues and the tracing of 
relationships would, of course, be much easier. We are, however, not 
without explanations for such variations as do occur. Fusions of chromo- 


C. E. MCCLUNG 


*72 

somes lessen the type number; splittings, fragmentations, non-disjunctions, 
supernumeraries, and multiple complexes increase it. Fusions have been 
most often observed in germ cells and there the continuity of all the elements 
has definitely been determined. Increases by various means are noted 
most commonly in somatic cells, but, as Hance has shown, there is probably 
neither loss nor gain of the chromatin. The importance of all these obser¬ 
vations lies in the convincing demonstration that numerical variations 
involve only rearrangements of the persistent chromatin aggregate. 

Of much significance regarding the evolution of chromosome groups are 
the observations upon mutating and hybrid species. While it is true that 
most of these have been made on plants their general value is just as great, 
because the fundamental facts of chromosome structure and behavior are 
common to all organisms. It appears here that changes in chromosome 
number result in differences of body structure. An additional element 
even, as in Oenothera scintillans, produces a different plant structure. 
When the group is duplicated, as in Oenothera gigas, all the parts of the 
plant are increased in size. Similar chromosome modifications do not 
accompany the mutations in Drosophila, however, and it has been suggested 
that the marked cellular instability in Oenothera is due to a hybrid character. 
Cytological support is lent to this view by the resemblances to similar 
conditions in known hybrids. Numerous observations have shown that 
when species with different numbers are crossed, during the maturation 
process there occurs a pairing between certain of the chromosomes, while 
others, unable to find mates, are cast out into the cytoplasm and disappear. 
In other cases they may persist and be carried along, dividing in one matura¬ 
tion division. At each maturation period of the hybrid generation rearrange¬ 
ments take place until a condition of stability is reached. Then, even 
in the presence of the normal number of chromosomes, a different type of 
organism results because of the change in the composition of the chromo¬ 
some complex. Studies upon cultivated plants show many instances of 
variations in chromosome numbers. Correlated with specific distinctions, 
Blackburn and Harrison found, for example, that in the genus Rosa the 
fundamental haploid number is seven and that there are certain species 
which are not only diploid but tetraploid, pentaploid and hexaploid. No 
forms with diploid complexes showed abnormalities in the maturation 
mitose, but in most tetraploid and in all higher multiple groups the number 
of chromosomes was reduced toward the type number. Similar results 
have been reached by Tahara in studies upon Chrysanthemum, byKihara 
upon wheat and by Kuwada upon corn. Sakamura finds that in the one 
grained wheats the chromosome number is 14 and that the multipled 


EVOLUTION OF THE CHROMOSOME COMPLEX 


73 


grained wheats show tetraploid and hexaploid groups. His suggestion 
that these represent an evolutionary series is in accordance with conclusions 
reached by other methods of reasoning. 

From the knowledge now before us it would, therefore, appear that the 
relation between the chromosome mechanism and the development of 
the characters of the species in the individual is very direct, in fact is a 
causal one. It is also apparent that the constancy of organization of the 
ultimate chromatin units into chromosomes is variable in different groups. 
In some, as in the Acrididae, this is very definite and fixed throughout the 
family, in other cases, e.g., the Hemiptera, it is more variable. With 
sufficient evidence it would, however, appear that there is always a strong 
tendency for the establishment of a number of chromosomes typical of the 
group, even in so large a one as a Phylum. The persistence of individual 
chromosomes in a foreign cytoplasm, making possible the hybridization of 
species, offers a means for changes in chromosome numbers. To some 
extent this is due to a differential reaction of the individual chromosomes, 
some apparently being more resistant or adaptable than others. The 
crucial test of chromosome compatibility is the synaptic period in the 
germ cells. Up to this point they may function with apparent normality, 
but when the chromosomes of the two parents are brought into this intimate 
union they often fail to survive the ordeal and imperfect germ cells result. 
Crosses between species with different chromosome numbers have fre¬ 
quently been made, however, and it is then found that distinct chromosome 
numbers accompanying differential body characters result. The means 
for an evolutionary change on the chromosome complex is thus afforded. 
Indeed our experience would indicate that to establish any change in 
hereditary characters, there must first occur a modification in the chromo¬ 
some complex, most commonly identified with a numerical alteration. 

Available evidence would also indicate that the chromosomes are made 
up of smaller units, each of which is in some respect functionally different 
from others in that chromosome, and from corresponding units in other 
chromosomes. It is not clear whether this differential nature in each 
case is due to an entirely unique activity or whether it is the result of a 
variability in degree, rate or time of action. Logically, and in consonance 
with our experience, it would be more reasonable to regard the factors 
as having much in common, with individual differences in form, degree, 
or time of action. In view of the great numbers of cell divisions required 
to produce an organism, each of which results in a slightly different charac¬ 
ter in the total constitution of the individual, it must be true that the extent 
of activity of any one factor is not great, a very slight difference in its 


74 


C. E. MCCLUNG 


operation each time being sufficient to produce an end result of larger value. 
Indeed it passes the range of human comprehension that there could exist 
a mechanism of such perfection as to have held a group like the Acrididae 
constant to type through such periods of time and through countless 
myriads of individuals. Differences in factors we must, therefore, assume 
to be of relatively minor nature. 

It would seem, moreover, that as between morphological and physiological 
distinctions among factors, we must conceive such to be largely physiological. 
Structural differences there surely are, but these are unlike those charac¬ 
teristics of form which we have found in units of higher order. Cells are 
distinguished apart by varieties of form, symmetry, polarity, proportion 
of parts, etc., all related in some way to similar distinctions between tissues 
and organs. The chromosomes themselves display some of these differences 
of a spatial nature, but they seem to bear no relation to the parts of the 
body. While nerve cells and muscle cells are distinctly unlike mor¬ 
phologically and arrive at such a state by gradual differentiation, the 
chromosomes of their cells show no corresponding form changes. It is 
true also that there are most extensive physical differences in the same 
chromosomes of a cell, due to its activities. 

Even more perhaps are distinctions between the ultimate chromatin 
units not those of what we call a morphological nature. For structural 
distinctions we should here, in all probability, have to seek those of a 
molecular character, and so would pass beyond criteria which we have 
employed in distinguishing between organisms and their parts. Physio¬ 
logically, however, it is possible to conceive of the same sorts of reactions 
carried down to the ultimate chromatin units. All physiological activities 
are essentially physico-chemical in their nature and a certain part of the 
body is, e.g. marked by a high development of some one of these, like 
irritability, contractility, or secretion. The determining factor here is in 
the activity of the nucleus and probably in the preponderant function of 
certain of the chromatin units. 

Taking all the available evidence into consideration, we must conclude 
that there is reasonable hope that we may be able to discover certain 
relations between the chromosomes of a group of organisms which will 
indicate to us their phylogeny. Very detailed studies, both cytological 
and genetical, should inform us of the specific nature of each chromosome. 
With sufficient of such group studies for comparison we may hope to reach 
some conclusions regarding the principles governing sub-division of func¬ 
tions between the members of a chromosome group. The test of homo- 
logues will largely be a functional one, structural differences serving as 


EVOLUTION OF THE CHROMOSOME COMPLEX 


75 


guides to identification. Work so far accomplished, which can only be re¬ 
garded as of the most preliminary character, is very encouraging and makes 
it reasonable to hope that not only will we arrive at a knowledge of the 
evolution of the chromosome complex itself but we will at the same time 
accomplish such an analysis of the constitution of one organism as to make 
clearer the relations which exist between different types. 


ABERRATIONS IN CHROMOSOMAL MATERIALS 1 

CALVIN B. BRIDGES 
Columbia University , New York City 

In Drosophila melanogaster a considerable number of distinct somatic 
types have arisen that have been shown to be the result of particular aber¬ 
rant groupings of chromosomes. For example, the mutant form “ Dimin¬ 
ished” is due to the loss of one member of the pair of fourth chromosomes 
(2n — IV). Also numerous aberrrations from the normal mode of inheri¬ 
tance have been traced to corresponding aberrations in the distribution of 
chromosomes or of sections of chromosomes. Most of our evidence as to 
the chromosomal conditions underlying these aberrations has been derived 
from genetical investigation, but the gentical finding has been successfully 
checked by microscopical examination in those cases in which the unit of 
change is a chromosome or the whole group of chromosomes (n). 

The various types of chromosomal aberrations and their effects may be 
included under the general term mutations, and may be called sectional, 
chromosomal, or n-mutations, according to the unit of change. Sectional 
and chromosomal mutations may be called ratio mutations, since their 
effects are produced by changes in the ratio in which genes are present, the 
chromosome being regarded as aggregates of genes. But these genes are 
themselves different, and new differences are apparently arising sporadically 
by transformation of old genes into new. Since the various effects seen in 
the ratio and n-mutations are referable to the action of component genes, 
all these types of mutation afford light on the characteristics of genes, and 
on how they interact to produce a given result. 

Analysis of the various mutational effects, and of the linkage relations, 
leads to the conception of a gene as a distinct chemical entity having a 
definite location in the chromosome. With each cell generation the original 
gene gives rise to two daughter genes identically located and (ordinarily) 
with the identical characteristics of the parental gene. In this increase in 
the amount of the substance of a particular gene, constituents that were 

1 Since this paper was read at the session of the Eugenics Congress it has been expanded 
and rearranged, and changed considerably in nomenclature. In making these changes 
other workers on Drosophila and on Datura have been consulted, but they are not com¬ 
mitted to any of the terms used here. 


76 


ABERRATIONS IN CHROMOSOMAL MATERIALS 


77 


present in the surrounding medium interact to produce a specific end 
product. 

To account for the occurrence of a mutant gene, it is assumed that the 
production of the old type of genic material has ceased and in its place a 
new type of material is being produced continually. In this new type of 
synthesis the raw materials present in the surrounding medium are essen¬ 
tially the same as those present in the previous situation. This is clearly 
seen in the fact that the cell may carry on both types of synthesis simultane¬ 
ously, as is the case in the heterozygote. The primary difference between 
the two types of synthesis to be sought then, not in the cytoplasm at large, 
but in the particular definite locus in which the synthesis occurs. Effective 
synthesis is thus seen to depend upon the action of some material situated 
at that particular locus; but when the synthesis has occurred the substance 
has not been used up, or else it has been reconstituted, for in the daughter 
loci the same reaction occurs at the succeeding growth period. This suggests 
that the reaction is catalytic, and specifically catalytic, since in the heterozy¬ 
gote one locus synthesizes one type of gene and the homologous locus 
synthesizes another. The situation can be accounted for on the assumption 
that the specific catalyst is the genic material itself; and this assumption 
is not out of harmony with the chemical conception of “ autocatalysis. ” 
The reaction may be called autocatylytic if the genic material is both 
catalyst and an end product. The molecules of the gene are assumed to be 
such that if the proper raw materials are present in the surrounding medium 
interaction occurs and one of the reaction products is the initial type of 
genic material, which thus increases in amount. 

The autocatalytic synthesis of a gene may be formulated: G -f- Cg —> 
2G + Pg, in which Cg represents the raw materials taken from the medium 
in the synthesis of G, and Pg represents the series of end products that are 
returned to the cytoplasm. If, as seems most probable, the genes act also 
as heterocatalytic enzymes, other specific end-products of greater variety 
are synthesized. Considered from the standpoint of the end-products, 
genes are chemical factories each of which is synthesizing a characteristic 
set of products which are delivered to the common cytoplasm. The various 
products influence the constitution of the cytoplasm directly by their 
presence; they may interact with one another, as well as with materials 
from outside; and they may act as raw materials in the syntheses carried 
out by other genes. 

The cessation of the synthesis of the old type of gene and the inception 
of the synthesis of a new form are accounted for by assuming that a chemical 
change has occurred in the genic material. (Go —> Gi . . . . Gi). 


78 


CALVIN B. BRIDGES 


The autocatalytic reaction Gi + C G i 2Gi + Pgi (together with whatever 
heterocatalytic action is present) liberates a set of end-products that is, or 
may be, different from that characteristic of the action of Go, and accord¬ 
ingly the diversity of cytoplasm and of development in characters proceeds 
step by step with the diversity of genes by mutation. Such transforming 
genic mutation (“ transgenation ”) is probably the most characteristic 
process in the production of mutant changes. 

In producing development the genes of the entire complement act 
together, since all are liberating their chemical products into the common 
cytoplasm. But since the products of the different genes are different they 
take effect in different ways upon the developing organism. Thus some of 
the genes will have much effect upon one character but little effect upon 
another. Each character will be determined by all the genes, but in each 
case most of the effect will be produced by a much more limited number. 
Each particular character shows a grade of development corresponding to 
the number and effectiveness of the genes that are producing it. In the 
case of many of the characters there are genes tending to make the character 
less pronounced, and the grade realized is that corresponding to the equilib¬ 
rium between these plus and minus modifiers. The directions of modifica¬ 
tion for each character are in general quite diverse, and it is somewhat 
for the sake of simplicity that they are lumped together as plus and minus 
modifiers. For example, the trident pattern on the thorax of Drosophila 
melanogaster may become darker or lighter in intensity, blacker or yel¬ 
lower in color, sharper or more diffuse, larger or smaller, and may change 
in shape by changes in any of its regions. While it seems probable from 
the facts of multiple allelomorphism that all the mutant forms of the genic 
material at a given locus have a like tendency, yet the situation is more 
complex, as the existence of specific and disproportionate modifiers shows. 
In genic mutation the point of equilibrium is shifted by the transformation 
of a modifier of one grade into a modifier of a different grade. 

It is observed that the presence of an extra chromosome (2n + 1) or the 
loss of a chromosome (2n — 1) causes character changes in the individual 
possessing the abnormal group. Such character changes can be interpreted 
in terms of component genes as follows: Linkage experiments show that the 
various kinds of genes are distributed indiscriminately among the various 
chromosomes and along each chromosome. But since the number of each 
kind is limited, short sections of equal length would not ordinarily contain 
the same proportion of any one kind, and still less would they contain the 
same proportion of every kind. The loss of one of these sections (a condi¬ 
tion known as “deficiency”) might remove more plus than minus modifiers 


ABERRATIONS IN CHROMOSOMAL MATERIALS 


79 


(or vice versa), and since relatively more minus than plus modifiers would 
then remain in action, the grade of the corresponding characters would be 
shifted in the minus direction. This is the interpretation of the fact that a 
deficiency may cause many character changes, this complex of new charac¬ 
ters being inherited as a simple dominant. In the case of the two dominant 
mutants Delta and Hairless, which show characteristics indicating that 
they are due to deficiencies, each presents a great variety of characters 
which are mainly opposite in direction in the two complexes. For example, 
the veins of Delta are heavy, and spread out at the marginal vein into deltas, 
while the viens of Hairless are delicate and vanish entirely at the margins. 
When these two types are crossed together the double mutant type is 
practically wild-type. The neutralization of these two dominants may be 
interpreted by supposing that Delta represents the deficiency of a set of 
minus modifires, that Hairless represents the deficiency of a corresponding 
set of plus modifiers, while the dihybrid has the original equilibrium re¬ 
stored. The gain of a section of chromosome (a condition called “ duplica¬ 
tion”) would likewise lead to character changes through changes in the 
ratio of genes. 

In a whole chromosome the ratios of plus to minus modifiers ought to be 
on chance closer to the ratios present in the whole complement, but on the 
other hand more kinds of characters might be involved, so that the character 
changes consequent upon the gain of a particular chromosome (through 
non-disjunction) might be both complex and extensive. 

The loss of a chromosome seems to produce greater character changes 
than does the gain of that same chromosome. Thus in Drosophilia melano- 
gaster individuals that have lost one of the pair of fourth chromosomes 
(2n — IV) through non-disjunction show many and extreme departures 
from the wild-type, while individuals that have gained a fourth chromo¬ 
some (2n + IV) show slight differences from the wild-type. The dif¬ 
ferences that are present in the extra-IV individuals are the opposite of 
those present in the minus-IV’s. Thus the minus-IV’s have large roughish 
eyes, pale body-color, dark trident-pattern, short blunt wings, etc., while the 
plus-IV’s have small smooth eyes, dark body-color, inconspicuous trident- 
pattern, long narrow wings, etc. This difference in sign between the charac¬ 
ters of the plus- and the minus-IV’s was expected on the basis of genic 
balance. From the genetical data it was evident that extra-IV individuals 
were present as half the flies in certain cultures, but they were not identified 
until a deliberate search was made for flies with characters the opposite of 
those already known for the minus-IV’s. In Oenothera plus but not minus 
deviants from the 2n condition are known. In Datura all the possible 


80 


CALVIN B. BRIDGES 


2n + 1 forms are known but none of the 2n— 1 forms have been found. 
These facts are interpeted to mean that the 2n— 1 forms are such extreme 
departures from the type that they are unable to survive. 

If the loss of a chromosome produces greater character changes than the 
gain of the corresponding chromosome then deficiencies would be expected 
to cause greater changes than corresponding duplications. There is some 
evidence that this is true. In the case known as “Pale-translocation” the 
genetical data lead to the hypothesis that the end of one of the second chro¬ 
mosomes has been broken off and joined to the third chromosome. By 
outcrossing this type to normal flies it is found that flies possessing the 
extra section (2n + Pale-section) are viable and not greatly different in 
characters from normal. But the flies that lack this section (2n — Pale- 
section) are so changed as to be entirely unable to live. In all the cases in 
Drosophila to which the hypothesis of deficiency is applied the “homozy¬ 
gous deficients” (2n — 2 section) are lethal, while in two of the three 
cases in which duplication effects are present the “homozygous duplicants” 
(2n + 2 section) are viable. 

If minus deviations tend to cause more extreme and more lethal character 
changes than do plus deviations, then deficiencies and the n — 1 gametes 
produced by non-disjunction should play a less direct role in evolution than 
do duplications and the n + 1 gametes. The plus deviations have the 
further possibility of being the basis for fresh diversity by transforming 
genic mutations. Presumably transgenation would go on independently in 
the (three or) four representatives of each of the genes of the unit involved. 
The identity and even the similarity of effect might be lost through large or 
successive transgenations. 

It has been a paradox that forms triploid or tetraploid (2n -f- n; 2n + 2n) 
are less inviable and less extreme in character changes than forms that are 
triploid for a single chromosome (2n +1). In Drosophila melanogaster, 
for example, the triploid (2n + n) differs from the wild-type only in certain 
size features; but the plus-IFs (2n + II) and the plus-ill’s (2n + III) are 
entirely unable to live, while the plus-X’s (2n + X) live only rarely and 
weakly. The relative similarity in characters of the 3n and 4n forms to the 
2n is understandable in terms of genic balance; for in these three forms 
the different genes are in identically the same ratios, while in the case of 
the 2n + 1 and 2n forms many of the ratios should be different. The 
differences that are present in the 3n and 4n forms may be due in part to 
the changed physical relations arising from the greater nuclear content, 
and in part from changed chemical relations connected with critical 
concentrations. 


ABERRATIONS IN CHROMOSOMAL MATERIALS 


81 


The origin of 3n and 4n forms is traceable to a division of the chromo¬ 
somes of a cell without division of the nucleus. This process is a kind of 
mass non-disjunction for which the term nuclear non-division (or simply 
non-division) may be convenient. Non-division, and indeed most of the 
mutational processes, may occur in the soma, giving rise to mosaics. Or 
they may arise early enough in development so as to involve both somatic 
and germinal tissue. This latter case should be especially frequent in 
plants. Self-fertilization following non-division, as in the case of a plant, 
would give rise to a tetraploid line; but in the case of animals, where cross¬ 
fertilization is the rule, the usual result of non-division would be triploid 
individuals. 

Besides the mutational processes mentioned, there are several others, 
some of which are included in the classification of the table below. The 
primary classification of mutation is made on the basis of the unit of change, 
and the secondary is according to the kind of change that can affect each 

unit. 

Mutation 


UNIT 

PROCESS 

RESULT 

Gene. 

Transgenation (Go—Kb* • *Gi) 

Deficiency 

Duplication 

Translocation 

Inversion 


Section.j 

n — 1 section 1 

. , .. ) gamete 

n + 1 section J 

New linkage relation 

New linkage relation 

Chromosome. 

Non-disjunction 

Elimination 

n + 1, n —1 \ 

. >gamete 

n 1 J 

n —> n r , where n r < n 

n — n', where n'> n 


Compounding 

Fragmentation 

Group (n) nucleus.j 

Nuclear non-division 
Nuclear fusion 

Binucleation 

Dispermy 

2n\ 

>gametes 

Mosaics 

Mosaics 













VARIATIONS IN THE JIMSON WEED (DATURA STRAMONIUM) 
CAUSED BY DIFFERENCES IN THE NUMBER OF 
CHROMOSOMES 

A. F. BLAKESLEE 

Carnegie Institution of Washington, Cold Spring Harbor , New York 

In normal (diploid) plants there are 12 pairs of chromosomes ora total of 
24 in each vegetative cell. 

If there is a single extra chromosome in a definite one of the 12 sets 
(making a total of 25), the plant shows specific peculiarities of structure. 
Thus, if the single extra chromosome is present in the chromosomal set 
which carries the determiners for the Mendelian characters purple or white 
flower color, the plant has, among other peculiarities, the arrangement of 
leaves which suggested the name Poinsettia for this mutant; if the single 
extra chromosome is in the set which carries the determiners for the presence 
or absence of spines on the capsules, the capsule is narrow suggesting the 
fruit of the Cocklebur for which the mutant has been named. An extra 
chromosome in a third set makes the plant a Globe mutant with globose 
capsules. Theoretically there are at least 12 “ simple trisomic” mutants 
possible with 25 chromosomes, each having 3 chromosomes in one particular 
chromosomal set and 2 in all the remaining 11 sets. 

The chromosomes contain factors or determiners for all the structures 
in the plant. One chromosomal set may contain determiners which tend 
to make the capsule long and narrow; another set may contain determiners 
which tend to make the capsule short and thick; and similarly for the leaves 
and other parts of the plant. The structure of the normal (diploid) plant 
is due to the resultant balance between the more or less opposing tendencies 
of the 12 chromosomal sets. In the simple trisomic mutants with a single 
extra chromosome in a specific set, the balance is disturbed and the deter¬ 
miners in this set have an increased influence upon the make up of the plant 
producing such mutants as Poinsettia, Cocklebur, and Globe. 

Plants may have 1 extra chromosome in each of the 12 sets as in “Tri- 
ploids” or 2 extra chromosomes in each set as in “ Tetraploids, ” but in 
such plants the influence of the several sets is balanced and the plants are 
not greatly different in appearance from normals (diploids). 


82 


VARIATIONS IN THE JIMSON WEED 


83 


Table 1 gives the chromosomal types already observed. 

TABLE 1 

Chromosomal types of mutants in the Jimson Weed (Datura Stramonium'). 
The somatic number of chromosomes is given in parentheses after each type. 


Balanced Types (12n) 

Diploids (Normal) (24) 

Triploids (36) 

Tetraploids (48) 

Unbalanced Types (12n + x) 

Modified Diploids (24 + x) 

A. Simple Trisomic (24 + 1) 

1. Globe 

2. Poinsettia 

3. Cocklebur 

4. Ilex 

5. Mutilated 

6. Sugar loaf 

7. Rolled 

8. Reduced 

9. Buckling 

10. Glossy 

11. Microcarpic 

12. Spinach 

Modified Tetraploids (48 + x) 

A. Simple Pentasomic (48 + 1) 

1. Globe 

2. Poinsettia 

3. Cocklebur 

4. Ilex 

5. Rolled 

6. Reduced 

7. Glossy 

8. Microcarpic 


B. Simple Tetrasomic (24 + 2, the 2 ex¬ 

tras in the same set) 

1. Round-leaf Globe 

C. Double Trisomic (24 + 2, one extra 

in each of 2 different sets) 

1. ex-Buckling (?) and a num¬ 
ber of other double mutants not 
yet analyzed by breeding tests 


B. Simple Hexasomic (48 + 2, the 2 ex¬ 

tras in the same set) 

1. Globe 

C. Simple Trisomic (48 — 1, a single 

deficiency in one set) 


Simple trisomic mutants produce normals (diploids) and similar simple 
trisomic mutants in their offspring; diploid and tetraploid plants breed 
true, while triploids produce normals and the different trisomic mutants. 




THE ATTRACTION BETWEEN HOMOLOGOUS CHROMOSOMES 


JOHN BELLING 

Carnegie Institution of Washington, Cold Spring Harbor, New York 

My thesis is very simple. In two plants, Canna and Datura, a mono¬ 
cotyledon and a dicotyledon, homologous chromosomes, when more than 
two in number, are attached to one another before the first maturation 
division, in threes, fours, or fives. 1 Thus the attraction between homologous 
chromosomes, which is the foundation of their conjugation in pairs and of 
their random assortment, and so of Mendelian ratios and of linkage dif¬ 
ferences, may extend to five (or six) such chromosomes. 

In place of cutting sections, pollen-mother-cells were squeezed from the 
anther into a mixture of 45 per cent acetic acid as fixative, ferric acetate 
as mordant, and carmine as stain. By varying the relative proportions, 
preparations could be obtained in which the chromosomes were nearly 
black, while the cytoplasm was unstained. After a day in the mixture the 
cytoplasm became plastic, and local pressure on the thin cover-glass broke 
the cell wall, allowing the contents to escape in a circular form while sufficient 
further pressure made the separate chromosomes assume a horizontal 
position. Among many such squeezes of single cells, some first prophases 
and second metaphases were found in which each chromosome or aggrega¬ 
tion of chromosomes was horizontal and sufficiently apart from its neighbors. 
(The work on Datura was all done with the cultures of Dr. A. F. Blakeslee.) 

In the normal Datura stramonium, the late first prophase shows 12 pairs 
of chromosomes, each chromosome usually transversely constricted, and 
each bivalent forming a ring, a V, or a bent or straight rod. Six size classes 
can be distinguished among the 12 bivalents (1 largest, 4 large, 3 medium, 
2 small medium, 1 small, and 1 smallest). The second metaphase shows two 
plates of 12 chromosomes each, every chromosome being divided into two 
transversely constricted halves, so that it appears four-lobed. The six 
size classes can be made out in each plate. (In a hundred cases, there have 
been only one or two instances of an assortment of 13 to 11, or of 12 to 1 
to 11.) 

1 This has lately been shown to be probab ly the case with six homologous chromosomes, 
also. (Note of October 28.) 


84 


ATTRACTION BETWEEN HOMOLOGOUS CHROMOSOMES 


85 


In Daturas with 25 somatic chromosomes, the prophases show 11 biva¬ 
lents like those of the normal plant; and also one trivalent, whose three 
chromosomes appear as a ring with attached rod, as a three-rayed aggre¬ 
gate, or as a bent rod. These three chromsomes are equal, and belong in 
different Daturas to different size classes. The second metaphase plates 
are usually 13 and 12, and in the plate with 13 an extra chromosome is found 
in one of the six size classes. 

In Daturas with two different chromosomes duplicated, we find 10 
bivalents in the prophase, along with two trivalents. The usual assortment 
is either 13 to 13, or 14 to 12, with about equal frequencies. The two extra 
chromosomes can sometimes be seen to belong to different size classes. 

In the prophases of triploid Daturas there are 12 trivalents, or aggrega¬ 
tions of three chromosomes each, these three chromosomes being of equal 
size. The six size classes can be readily distinguished. In the second 
metaphase the assortment may be seen to vary from 12 and 24 through all 
the intermediates to 18 and 18. The numbers accord fairly well with the 
binomial distribution expected if the trivalents are oriented at random on the 
spindle. In some buds there are many, and in others few cases of isolated 
chromosomes not included in either second metaphase plate. 

In tetraploid Daturas the chromosomes are in fours in the prophase of 
the first division, forming quadrivalents or bigemini (Marchal). These 
appear usually as double rings, or as crosses. Usually two chromosomes 
of each quadrivalent go to each pole, but it is probable that sometimes three 
go to one pole and one to the other. The six size classes can be readily 
distinguished, and the four chromosomes of each quadrivalent are of the 
same size. The assortment is usually 24 to 24, there being two chromosomes 
of each kind in each metaphase plate; but not uncommonly 23 to 25 is 
found, or even 22 to 26. 

In tetraploid Daturas with one extra chromosome, the assortment is 
usually 25 to 24, and in the second metaphase plate with 25 chromosomes 
there is one extra chromosome in one of the size classes. In the first pro¬ 
phase a quinquevalent of five chromosomes has been twice seen. 

One tetraploid Datura with two extra chromosomes was examined. 
The common assortments were 25 to 25, and 26 to 24. In the second 
metaphase plate with 26 chromosomes, one of the size classes which usually 
shows here only two members in tetraploid plants occurred four times, 
while it was found twice in the corresponding 24-chromosome plate. Thus 
this chromosome occurred six times in the first prophase. (Two prophases 
of this plant have lately been examined, and each shows only 12 aggregates. 
It is probable that one of these is a sexivalent. Note of October 28.) 


LINKAGE WITH LETHAL FACTORS THE SOLUTION OF THE 
OENOTHERA PROBLEM 

GEORGE H. SHULL 

Princeton University, Princeton , New Jersey 
INTRODUCTION 

There is perhaps no better way to open a paper on this subject than by 
quoting the following statement from Muller’s paper on “An Oenothera- 
like case in Drosophila,” which appeared in the Proceedings of the National 
Academy of Sciences for October, 1917: 

In Oenothera a form of the balanced lethal explanation was suggested by de Vries, only 
to account for his double reciprocal crosses, but it is evident from the analogy of the 
beaded case that it probably lies at the root of nearly all the unusual genetic phenomena 
of the genus. The two cases differ in detail, however, in that one or more of the lethals 
in Oenothera produce their effect upon the gametes, rather than upon the zygotes. 

Beginning in 1913 and proceeding step by step I have been able to demon¬ 
strate the truth of Muller’s statement to such an extent that the title 
of my paper seems fully justified by the data which have been secured. 
It will be impossible in the short time at my disposal here to do more than 
sketch the mechanism of heredity in the Oenotheras as it now stands clearly 
revealed, and to present samples of the evidence which has been secured. 

In brief, the elements of the genetic machine which is operating in the 
Oenotheras, are as follows: There are seven pairs of chromosomes which, 
according to many careful cytological studies, are typically behaving during 
gametogenesis and fertilization just as do the chromosomes of other species 
of'plants and animals. There is thus provided a mechanism capable of 
producing in Oenothera all of the genetical phenomena which are attribut¬ 
able to chromosome distributions in other organisms. 1 The “Oenothera 

1 1 refer here only to the orderly succession of diploid and haploid generations, with 
the certainty that homologous maternal and paternal chromosomes are separated into 
different germ cells. In a recent note (Jour. Heredity 13: 75, 76, 1922) Gates calls 
attention to the well known fact that the chromosomes in Oenothera “form a chain end- 
to-end like a string of sausages, and when they ultimately come to be side-by-side in 
diakinesis they are already in the short and stout condition in which twisting about each 
other is impossible.” He thinks this fact must be “very disturbing” for those who write 
about crossing over in Oenothera. The empirical genetical evidence for crossing over 

86 


SOLUTION OF THE OENOTHERA PROBLEM 


87 


problem” may be formulated, then, in a sentence, thus: Why is it that with 
chromosomes visibly behaving as they do in other organisms, extensive 
and intensive breeding experiments covering nearly thirty-five years have 
led many able geneticists to the view that Oenothera is genetically “a law 
unto itself?” The answer to this question is as follows: With the exception 
of one character (the short style of brevistylis) all of the characteristics 
thus far investigated, by which the numerous Oenothera species are differ¬ 
entiated, are determined by factors located in a single pair of chromosomes, 
which I have designated “chromosome I,” and the evolution of the genus 
Oenothera, in so far as it relates to these characters, reduces practically to 
the evolution of this single pair of chromosomes. 2 One of the first steps in 
this evolution was the origin of one or more recessive lethal factors, whose 
presence has acted like a mask behind which other genetic changes have 
taken place without being subjected to summary elimination in the struggle 
for existence. The evolutionary significance of the lethal factors appears 
to be not alone, however, that of a protection to other mutative changes, 
which Muller speaks of as chromosome degeneration; for a compari¬ 
son of forms having the lethals, with corresponding forms which do not 
have them, indicates that the lethals may have had a positive r6le in the 
evolution of the group in addition to their passive r61e in protecting other 
factors from elimination. The alethal forms are usually inferior in vegeta¬ 
tive vigor, or rapidity of development, or degree of fecundity, as compared 
with the lethal-bearing types, so that it seems probable that the lethals 
are either themselves stimulative in effect or else that they act as preservers 
of the stimulating effect arising from the interaction of the unlike factors 

in the Oenotheras is unequivocal, however, as indicated in a preliminary way in this 
paper, and this evidence has been greatly strengthened and extended since this paper was 
written. It is only fair to point out in this connection, that genetical analysis of chromo¬ 
some behaviors is vastly more refined and effective than can be attained by present 
cytological technique, so that if there is a real antagonism between the evidence from the 
two methods of investigation, the cytological studies are more likely than the genetical 
to be in error through not having yet attained their proper goal. 

* O. Renner (Versuche fiber die gametische Konstitution der Onotheren. Zeitschr. 
indukt. Abstamm.- u. Vererb. 18: 121-294. 1917.) has given to a number of the gene- 

complexes which constitute the two members of chromosome I in various species, distinc¬ 
tive Latin names; but he understands by these names the genotype contained in the 
entire group of seven haploid chromosomes instead of the linkage-groups which constitute 
the single chromosomes of one pair, as they now appear to be (see op. cit., p. 229). Thus, 
the two gene-complexes which constitute the two members of this pair in Oenothera 
Lamarckiana are called “velans” and “gaudens”; the two in Oenothera biennis are 
“albicans” and “rubens”; the two in Oe. suaveolens are “albicans” and “flavens”; and 
the two in Oe. syrticola are “rigens” and “curvans.” 


38 


GEORGE H. SHULL 


(or the additive effect of dominant factors (Jones)) lying in the two mem¬ 
bers of chromosome I, which effect would seem to be analogous if not 
identical with the heterosis so frequently observed in the hybrids of other 
organisms. 

The early origin of the lethals and their pronounced effect on the vigor 
and fecundity of their possessors account for their very general occurrence 
in the different species of Oenothera, and it is their general presence which 
has thus far limited the occurrence of those types of genetic phenomena with 
which we have become familiar in other organisms and whose relation to 
chromosome-behaviors has been fully established by a number of critical 
lines of evidence. 

The lethal factors of Oenothera are of two kinds, the one kind destroying 
the zygote, the other either destroying one or the other class of gametes, or 
at least preventing them from functioning. It is the latter class of lethals— 
the gamete lethals—that are peculiarly characteristic of the Oenotheras, 
and which have been indicated as yet in only a few other plants and so far 
as I am aware have not yet been noted in any animal. 

With this brief general statement of the conclusions to which my investi¬ 
gations have led me, we are now prepared to consider the nature of the evi¬ 
dence, which has been consistently presented, in repeated tests, often in a 
large number of matings, and for which there seems to be no other satis¬ 
factory interpretation than the one here given. I believe that, taken in the 
aggregate, these data amount to a demonstration. 

It may be well to point out that, had the Drosophila breeders been con¬ 
fronted with the beaded-wings situation, not as an exceptional case which 
could be left on one side until a background of experimental evidence had 
accumulated from simpler situations which could be used with confidence as 
a basis of interpretation, but as the general type of nearly all Drosophila 
genetics, there would probably have been no more rapid progress in the 
unraveling of the Drosophila mysteries than there has been in the case of 
Oenothera. 

Just as it was necessary in Drosophila to have the simpler situations 
worked out, in order later to be able to control the elements of a situation so 
complex as that of the beaded wings, so in the case of Oenothera, it has been 
needful first to find clear cases of linkage of factors without the confusing 
effect of lethals, before any confidence could be felt in an interpretation of 
the more involved cases in which the masking effects of lethals were present. 


SOLUTION OF THE OENOTHERA PROBLEM 


89 


EXPERIMENTAL EVIDENCE 
Linkage between rubricalyx buds and red stems 

The first two cases which met this requirement came under observation in 
1913, but were held with suspended judgment or even in a skeptical frame 
of mind until 1915 and 1916, respectively, when they were considered as 
clearly established cases of linkage in the sense in which we have become 
familiar with it in the Drosophila literature. Both of these linkages 
involved the rubricalyx pigmentation of the buds (. R h ) in which the whole 
bud, except the ovary and the free tips of the calyx, are intensely reddened. 

Green stems, 

Red stems, green hypanthia 

red hypanthia (red cones) 

R* ♦ R h X r c . R* 



Red stems, green hypanthia Green stems, red hypanthia 

Fig. 1. Diagram showing the recombinations of the two chromosomes of pair I in 
a cross between red-stemmed rubricalyx and a form with green stems and green hypanthia 
like erythrina. The symbols in this diagram are to be understood as follows: R h f a 
dominant factor for red hypanthium, i.e., rubricalyx; R a , a dominant factor for intensely 
reddened stems; R c , a factor for red bud-cones, associated with recessive green hypanthia; 
r c , a recessive factor for wholly green buds. 

The other two factors which were at this early date found to be linked 
with the factor for rubricalyx pigmentation were the factor n for the dwarf 
stature of nanella, and R s which produces an intense reddening of the stems. 
The relation between red buds and red stems was such as to indicate that 
the corresponding factors, R k and R s , are allelomorphic to each other (or 
at least nearly so), for every plant tested, in which both red stems and 
rubricalyx buds were found, produced germ cells of two kinds only, with 
respect to these factors, namely, (a) those carrying the rubricalyx factor, 
R h , and (b) those containing the red-stem factor, R s . This discovery served 






90 


GEORGE H. SHULL 


to clear up immediately the peculiar negative correlation which I had pub¬ 
lished in 1914, for with such a constitution, a cross between a red-stemmed 
rubricalyx (R'R ft ) and a green-stemmed plant with green hypanthia (r c R c ) 
such as Lamarckiana or erythrina, gives a case of criss-cross inheritance of 
the type made familiar in cases of sex-linked inheritance. 

This situation may be shown simply by a diagram, as in figure 1. 

Self-fertilizing a plant of the red-stemmed rubricalyx type, R s . R h , results 
in a segregation of the Andalusian type, as shown diagrammatically in 
figure 2. 

This is the situation described in my 1914 paper as a “peculiar negative 
correlation.” 

Red stems, red hypanthia X self 


r* • R h X R' • R h 



Dark red stems, Bright red stems, Green stems, 

green hypanthia red hypanthia red hypanthia 

Fig. 2. Recombinations of the two chromosomes of pair I when a red-stemmed rubri¬ 
calyx plant is self-fertilized. Symbols as in figure 1. 

Ped. No. 1468, grown in 1915, was a small family showing approximately 
the expectation in this case, having 9 red-stemmed green-budded : 18 
red-stemmed rubricalyx: 6 green-stemmed rubricalyx, and a larger family 
grown in 1916 (Ped. No. 1592) gave the same groups in the ratio 40:84:42. 
No red-stemmed rubricalyx has been tested which did not, when selfed, 
segregate in the manner indicated. 3 

3 While Gates has repeatedly described his rubricalyx as having notably red stems, and 
I, too, have observed that its stems are more strongly reddened than those of rubrinervis 
or erythrina, its stems are still in what I call the green-stemmed group and in rather 
striking contrast to the intensely reddened stems produced by the factor, R*. In the 
autumn the green-stemmed group often redden up to an extent which makes classification 
difficult. 





SOLUTION OF THE OENOTHERA PROBLEM 


91 


Linkage between rubricalyx buds and nanella stature 

The other original case of linkage, of which I was at first skeptical, but 
which was accepted as such in 1916, through the observation in that year 
that the one infrequent crossover class, rubricalyx nanella, was balanced by 
an equally infrequent occurrence of the corresponding crossover class, 
green-budded tall, was that between the rubricalyx factor, R h , and the nanella 
factor, n. 

As an example of the evidence of the linkage of these two factors, I may 
cite the data from three F 2 families from a cross between rubricalyx tall 
and green-budded dwarfs. These three families totaled 213 rubricalyx 
tall, 1 rubricalyx nanella, 3 green-budded tall, 50 green-budded nanella, 
instead of 156:52:52:17 which should have been approximated had the 
two factor pairs been independent instead of being closely linked. 

A cross in which the two factors, R h for rubricalyx pigmentation, and n 
for nanella stature, came in from the same side of the parentage instead of 
from opposite sides as in the foregoing example, gave a corresponding result, 
but with the two middle terms preponderating, as is expected in crosses of 
this type. Five F 2 plants (tall rubricalyx) from this cross, were back-crossed 
to green-budded nanella and gave in 1920 the ratio 22 rubricalyx tall: 244 
pink-coned tall : 137 rubricalyx dwarfs : 11 pink-coned dwarfs. As 
the nanella plants are less easily reared than the tails there is some dif¬ 
ferential elimination of nanella plants, but as this seems to have affected about 
equally both the crossover class and the non-crossover class, the percentage 
of crossing over is not essentially modified. This back-cross test therefore 
suggests a crossover value between rubricalyx and nanella of about 9 units. 
The data are not yet adequate, however, to give this determination more 
than tentative value, as different tests have varied considerably in their 
indications as to the crossover percentage, though all agree qualitatively 
that the two factors in question are rather closely linked. 

Linkage between pink-coned buds and nanella stature 

The linkage between pink-coned buds, and nanella stature is indicated by 
an F 2 ratio from four selfed Fi plants, resulting from a cross between pink- 
coned tall and green-budded nanella. The F 2 of these four families consisted 
of a total of 206 tall with pink-coned buds, 20 tall with green buds, 11 nanella 
with pink cones, and 129 nanella with green buds. 

Linkage between sulfur-colored flowers and red stems 

In 1920 I had a large series of families representing a dihybrid combina¬ 
tion between sulfurea flower color and red stems. Five of the families were 


92 


GEORGE H. SHULL 


from selfed Fi plants and 11 from crosses between sibs in the Fi genera¬ 
tion. These 16 families gave consistent results, and aggregated 1149 red¬ 
stemmed yellow, 61 red-stemmed sulfur, 77 green-stemmed yellow, and 
686 green-stemmed sulfur. In this case I am tentatively assuming on the 
basis of other experiments that a lethal factor has cut down the class of red¬ 
stemmed yellows to about half their normal number and that the linkage 
value of 5 and R s is probably about 26 units. 

Linkage of revolute leaves with red stems and nanella stature 

In the present season (1921) I have had several families which tested the 
linkage relation between a factor, 4 /, for the revolute-margined leaves of a 
mutant type which I call funifolia, and the red-stem factor, R s , —also between 
revolute leaves and rubricalyx buds, R h . The form with revolute leaves 
first appeared in my cultures (Ped. No. 1720) in 1918, as the result of a gene 
mutation which probably occurred in 1916. It had green stems and pink- 
coned buds. The funifolia segregates in F 2 have had in both the crosses 
mentioned above only the green stems and pink bud-cones of the original 
specimens of this type or the red-coned buds of an erythrina mutant of 
funifolia which was used in several of the crosses. Not a single case of 
crossing over between revolute leaves and bud color has been observed. 5 6 
Thus, a cross between a green-budded, red-stemmed, flat-leaved dwarf 
(nanella) and a tall, green-stemmed, revolute-leaved funifolia, gave an Fi 
of all red-coned, red-stemmed, flat-leaved, tall plants, and an F 2 by self- 
fertilization, consisting of only three types instead of the 16 which would 
have been present had the four known alternative characters been inde¬ 
pendently inherited. These three F 2 phenotypes were the two parental 
types and the Fi type, the two former being presumably homozygous and 
the latter heterozygous, thus again presenting the Andalusian type. The 
lack of crossovers between / and R c is probably due to the inadequate num¬ 
bers of individuals in these tests. 

Linkage of revolute leaves with rubricalyx buds 

The F 2 from a cross between funifolia, which had long styles as in La- 
marckiana, and a rubricalyx brevistylis, gave a similar result. In this cross 
both of the chosen parents had tall green stems, so that only three alterna- 

4 For description and figures of Oe. mut. funifolia, see Jour. Heredity 12: 354-363. 

1921. 

6 Crossovers have appeared in 1922, giving both rubricalyx-budded funifolia and red¬ 
stemmed funifolia. 


SOLUTION OF THE OENOTHERA PROBLEM 


93 


tive characters were involved, namely, flat vs. revolute leaves, red vs. 
green hypanthia, and long vs. short styles. 

The Fi was uniformly flat-leaved, long-styled rubricalyx, giving on the 
basis of independent inheritance, an expectation of 8 phenotypes in F 2 , 
but the F 2 consisted of only 4 phenotypes, instead of the 8 expected, the two 
new types being due to the independent assorting of the brevistylis charac¬ 
ter with either of the other parental combinations. All of the funifolia 
plants had red-coned buds with green hypanthia and all of the flat-leaved 
plants had rubricalyx buds,—intensely red on both cones and hypanthia. 
Brevistylis plants were found in approximately the expected Mendelian 
proportions in both the funifolia and the rubricalyx groups. This family, 
then, shows clearly the same relation between revolute leaves and rubricalyx 
buds, that the previously mentioned family demonstrated between revolute 
leaves and red stems. This is in full accord with expectation, in view of the 
fact that rubricalyx buds and red stems had been demonstrated to be allelo¬ 
morphic to each other. If crossing over is possible between any two of these 
three factors it is certainly of very low frequency, and for the present it is 
held, tentatively, that the four factors for rubricalyx buds ( R h ), red stems 
(R s ), red bud-cones (. R c ) and green buds ( r c ) form a system of quadruple 
allelomorphs, and that the factor for revolute leaves (f) occupies a nearby 
locus. 

Having thus demonstrated that these five factors occupy the same (or 
very nearly the same) locus in chromosome I and that the factors for tall 
vs. nanella stature lie only a short distance from these, while the factors 
for yellow vs. sulfurea flowers are considerably more remote though still 
very obviously linked with them, let us turn to the characteristic lethals. 

The characteristic zygote and gamete lethals 

The existence of the gamete lethals was held tentatively as a working 
hypothesis for a number of years by Oenothera students in connection with 
the phenomenon of heterogametism which manifested itself in the pro¬ 
duction of unlike reciprocal hybrids. The presence of a pair of zygote 
lethals in Oenothera Lamarckiana also came to be strongly suspected be¬ 
cause of the general occurrence of segregation in the Fi generation of its 
crosses with other species and with many of its own derivatives. The 
difficulty, however, in demonstrating that Oe. Lamarckiana owed its 
peculiarity in this regard to the presence of lethal factors and that it was 
really a perpetual heterozygote in the Mendelian sense, lay in the fact that 
the Fi phenotypes seemed to differ from one another in almost every 


94 


GEORGE H. SHULL 


character instead of being differentiated from one another, as in the segre¬ 
gates of ordinary heterozygotes, by single characteristics. In fact, as I 
pointed out in 1914, the situation was so obscure that we could not at that 
time decide with any confidence, just what constituted a unit character 
in the Oenotheras. Two ideas, other than that of linkage with lethal 
factors, were favored by the complex nature of the segregates, namely, 
(a) the nuclear-chimera idea that chromosomes were hanging together in 
clusters, and (b) that there were so many non-compatible combinations of 
characters that only a small number, represented by the combinations which 
actually appeared, were capable of existence. 

Lamarckiana Rubricalyx 



bud color 

Fig. 3. Recombinations of chromosomes of pair I when Oenothera Lamarckiana 
with pink bud-cones and green hypanthia, is crossed with rubricalyx. Besides R h , R e > 
and which have the same significance as given under figure 1, h and k are two recessive 
zygote lethals, and L x and U are the dominant “normal” vital factors which are allelo¬ 
morphic to these lethals. 

Not until a known unit-factor could be followed through the genetic 
machinery was it possible to eliminate these seemingly plausible con¬ 
ceptions and establish in their place an interpretation which is in accord 
with the facts: (a) that cytological studies show no fundamental deviations 
in the chromosome mechanism, from that found in other organisms; and 
(b) that the numerous, very divergent species of Oenothera are quite 
generally fertile inter se, with only a moderate degree of sterility in the 
individual crosses. 




SOLUTION OF THE OENOTHERA PROBLEM 


95 


The zygote lethals of Oenothera Lamarckiana 

The demonstration that Oe. Lamar ckiana is really heterozygous in the 
Mendelian sense was made in 1917, with the analysis of a cross between 
Lamarckiana and rubricalyx. In order to make the situation easily com¬ 
prehensible, it is necessary to note that in Oe. Lamarckiana each member 
of chromosome I contains a recessive lethal factor, which we may desig¬ 
nate h and 4, respectively, while in Oe. rubricalyx, as also in the corre¬ 
sponding forms, Oe. rubrinervis and Oe. erythrina, only one of the chromo¬ 
somes contains a lethal, h or 4, while its mate has no zygote lethal. 
When a cross is made between Lamarckiana and homozygous rubricalyx, 
the Fi progeny splits into the two types, Lamarckiana and rubricalyx, with 
respect to all characters except color of the buds, which are uniformly 
rubricalyx throughout both of the Fi types, except that in the Lamarckiana 
group the pigmentation is usually a little less intense than in the rubricalyx 
group. This cross may be diagrammed as in figure 3. 

From this diagram it is seen that the expected ratio between Lamarckiana 
and rubricalyx plants in the Fi is 1:2. Leaving out of account three 
“mutant” forms in each of the two original reciprocal families from this 
cross (Ped. Nos. 1534 and 1535) the results secured in 1916 were: 

(rubricalyx X Lamarckiana) Fi 42 Lamarckiana : 100 rubricalyx 

(Lamarckiana X rubricalyx) Fi 25 Lamarckiana : 37 rubricalyx 

Total 67 Lamarckiana 137 rubricalyx 

Expected 68 Lamarckiana 136 rubricalyx 

In the F 2 , which was grown extensively in 1917, the rubricalyx type of 
F 2 plants gave splitting progenies in approximately the expected 2:1 ratio, 
10 families aggregating 863 rubricalyx: 326 with green hypanthia. The 
deficiency in the latter group was probably due to the fact that the homozy¬ 
gous form is less vigorous than the heterozygous. 

When the Fi plants having the vegetative characters of Lamarckiana 
and the bud pigmentation of rubricalyx were selfed, they gave, in 9 families 
representing progenies of plants from both of the reciprocal matings, 396 
plants having the form of Lamarckiana, or of several of its rare “muta¬ 
tions, ” and every plant had rubricalyx buds. In other words, although these 
plants had just been made heterozygous by the mating of gametes unlike 
with respect to the known factor for rubricalyx buds, there was no splitting 
out of the green-hypanthium character of the Lamarckiana grandparent. 

That these same Lamarckiana plants, which bred true with respect to the 
rubricalyx buds, were actually heterozygous with respect to the factor, 



96 


GEORGE H. SHULL 


R h , was demonstrated at the same time by back-crossing to Lamarckiana 
with its usual type of pink-coned green-hypanthium buds. The resulting 
progenies from 10 such back-crossed individuals aggregated 347 rubricalyx 
and 349 pink-coned or green-budded, all of Lamarckiana form except a 
few of the so-called “mutants.” This experiment at once established the 
correctness of the balanced-lethal hypothesis which had been up to that 
time held only tentatively as a working basis. 

It is to be noted that these balanced lethals, h, and 4, in Lamarckiana 
are zygote lethals. The microscopic evidence which is frequently cited in 
support of the balanced lethals, namely, that half the pollen and half the 
ovules fail to develop, would seem to be dealing with a wholly different set 
of lethals,—namely, gamete lethals. I know of no convincing evidence yet 
from genetic experiments that gamete lethals occur in Oe. Lamarckiana 
and its derivatives. The fact that the above-pictured results with the 
Lamarckiana-rubricalyx crosses gave the same ratios in the reciprocal 
matings bears against the existence of gamete lethals in chromosome I of 
this group. It is, of course, possible that gamete lethals occur in one of the 
other chromosomes, but in this case they can not distort the genetical 
results with respect to characters whose factors he in chromosome I. 

Demonstration of the gamete lethals 

The existence of gamete lethals as factors lying at definite loci in chromo¬ 
some I, has also been demonstrated in my cultures during the past two 
seasons, in two independent groups of material concerned with the problem 
of yellow vs. sulfur-colored flowers. De Vries has worked out the genetical 
behavior of this pair of alternative characters in both Oe. biennis 6 and Oe. 
suaveolens. 7 In both cases the color of the flowers in the hybrid families 
was typically that of the male parent. Thus S X Y = Y and Y X S = S, 
and if the Fi plants are selfed or back-crossed to the parent types, the F 2 prog¬ 
enies show no segregation, but again agree in flower color with the imme¬ 
diate male parent. This result is to be explained as follows: The wild types 
of both of these species carry a recessive factor, s, for sulfur-colored flowers 
in one member of chromosome-pair I, associated with a sperm-lethal, l s , while 
the mate to this chromosome carries the corresponding dominant factor for 
yellow flower-color associated with an egg-lethal, l e . 

6 De Vries, H., Gruppenweise Artbildung unter spezieller Beriicksichtigung der Gat- 
tung Oenothera, viii + 365 pp. Gebr. Borntraeger: Berlin. 1913. See pp. 297, 298. 

7 De Vries, H., Mutations of Oenothera suaveolens Desf. Genetics 3: 1-26. 1918. 
See pp. 8,9. 


SOLUTION OF THE OENOTHERA PROBLEM 


97 


When such a plant produces eggs, the 50 per cent which receive the yellow 
factor, S, are prevented from functioning by the egg-lethal with which this 
factor is linked, and the sperms, correlatively, which carry the factor for 
sulfur-colored flowers, 5, are prevented from functioning by the sperm- 
lethal linked with it. Thus it is that functional eggs carry only the factor 
for sulfur-colored flowers, while functional sperms have only the factor for 
yellow flower-color. 

The demonstration of the correctness of this picture has come through 
the occurrence of repeated crossovers between the gamete lethals and the 
factors for flower color. It is found that the association between the factor 
for yellow flower color and the egg-lethal and that between the factor for 
sulfur color and the sperm-lethal can be exactly reversed, so that the 
e gg-lethal goes with sulfur and the sperm-lethal with yellow, and it has 
also been demonstrated that in the absence of these lethals, typical 1:1 and 
3:1 ratios occur with respect to flower color. 

CONCLUSIONS 

With the foregoing evidence before us that the known genetical phenom¬ 
ena in the Oenotheras are almost wholly attributable to elements lying 
in chromosome I, let us enumerate some of the conclusions which appear 
to be justified. 

The Fi splitting and unlike reciprocal hybrids which have constituted 
the most fundamental phases of the “Oenothera problem” are specifically 
accounted for by the demonstrated zygote and gamete lethals. The great 
advance which is made by my studies in regard to these is that they are 
shown to be factors which occupy definite loci in a chromosome , and that we 
can appropriately speak of lethal factors ,—not merely “lethal gametes,” 
“lethal combinations of chromosomes,” or “lethal chromosomes.” 

A rational explanation is thus given for “ heterogametism ” and the so- 
called “ mass-mutations” of Bartlett and Miss Cobb, 8 since a “ single maternal 
or paternal chromosome” is substituted for “a distinctly maternal or dis¬ 
tinctly paternal set of chromosomes. ” “Mass-mutations” are to be under¬ 
stood as the result of crossing over of recessive genes lying at considerable 
distance from the masking lethals. The term is hardly appropriate as 
applied by de Vries to the simple 2 : 1 segregations which take place 
when only one member of chromosome-pair I carries a zygote lethal, as in 
the case of rubrinervis, erythrina, rubricalyx, etc. 

8 Cobb, Frieda, and Bartlett, H. H. On Mendelian inheritance in crosses between 
mass-mutating and non-mass-mutating strains of Oenothera pratincola. Jour. Washing¬ 
ton Acad. Sci. 9: 462-483. October 4, 1919. See p. 475. 


98 


GEORGE H. SHULL 


We are now in position to classify the “mutations” of Oenothera under 
three heads: (a) gene mutations, (b) crossover mutations, and (c) anomozy- 
gous mutations (i.e., chromosome aberrations). It is not proper to dis¬ 
criminate among these, as some writers have done, by designating (b) and 
(c) “de Vriesian mutations,” and (a) “Morgan’s mutations;” such a 
distinction is not fair to either of the men mentioned, nor is it fair to history. 
It is not possible at this late date to eliminate the term “mutation” from 
the literature of Oenothera as a convenient term by which to designate 
genotypically distinct forms which appear with a frequency low enough 
to be in striking contrast with the larger categories which make up the bulk 
of any given progeny. Only genetical experiments can distinguish between 
(a) and (b), and cytological evidence is required in the case of (c). 

The idea of “nuclear chimeras” is definitely set aside as unnecessary, and 
by the tracing of the major part of the sterility to definite lethal factors, 
Oenothera is made ineligible as an example of the phenomena depicted by 
Goodspeed and Clausen 9 under the name of a “reaction-system,” though it 
does not, of course, prevent each Oenothera genotype from being, like a 
genotype in any other organism, an example of a harmoniously operating 
reaction-system. 

All of the phenomena which have been attributed to a putative hybrid 
origin for Oe. Lamarckiana are accounted for, and although there is no 
ground for denying that Oe. Lamajrckiana, like any other species, may have 
had a hybrid origin, there is no longer any reason to lay stress on this possi¬ 
bility, since the genetical behavior of this species is shown to rest entirely upon 
its present factorial constitution. There is absolutely no evidence that a 
lethal factor or any other genetic factor can be brought into existence as a 
result of hybridization. 

The general compatibility of the various Oenothera species, notwithstand¬ 
ing their very striking divergences in character, is made easily compre¬ 
hensible when it is considered that these divergences are limited in great 
measure to the divergent constitutions of a single chromosome pair, thus 
leaving 12 of the 14 chromosomes with probably essentially identical con¬ 
stitution throughout the entire group of Oenothera species which have 
yet been studied genetically. 

Finally, the status of Oenothera as an object of research is materially 
improved, and the stone which has been rejected by certain of the builders 
is restored to its place as the “head of the corner,” for Oenothera becomes 
at once the plant which has the largest number of known factors lying at 

9 Univ. of California Publ. 5“: 301-322. Pis. 37-48. 1917. 


SOLUTION OF THE OENOTHERA PROBLEM 


99 


definite loci in a single chromosome pair or linkage group; the number 
of distinct hereditary characteristics available within the genus Oenothera 
is apparently almost limitless; and new gene mutations are taking place 
with encouraging frequency. While the general mechanism of heredity 
in Oenothera now stands revealed in a fair degree of completeness, it can 
not be too strongly emphasized that the details are only qualitatively correct, 
and that a very great amount of work remains to be done in determining 
the exact quantitative relations of the known genes to each other, and in 
adding new loci to the map of chromosome I, as well as in discovering factors 
which may possibly exist in the other chromosomes. The door stands wide 
open, and many students must join hands in the colossal task of exploring 
the recesses which lie within, in order to bring to light quickly the genetical 
treasures which are certainly there. 


A NEW TYPE OF VARIABILITY IN PLANTS 
R. RUGGLES GATES 
University of London (King’s College) 

The study of size-inheritance presents difficulties which do not appear 
with other forms of inheritance. These difficulties are chiefly due to (1) 
the absence of dominance, ( 2 ) obscuration of the results by the presence 
of fluctuations. The inheritance of nanism and gigantism is, however, 
fairly well understood. Many writers have explained size-inheritance 
where an intermediate Fi generation is obtained and a wide range of sizes 
in F 2 , as the result of the redistribution of several cumulative Mendelian 
factors for size. This conception has been applied particularly by East, 
Emerson, Hayes and others to the inheritance of such characters as length 
of cob in maize and length of corolla and size of leaf in tobacco. Certain 
recent facts make it necessary to re-examine more critically the whole matter 
of size-inheritance. It is possible that the inheritance of size in repeated 
parts such as flowers or leaves may follow a different law from the in¬ 
heritance of, for example, size or weight of the whole organism in crosses 
between large and small varieties. 

In crosses between large-flowered and small-flowered species of Oenothera, 
the Fi is intermediate in flower size, while it is found that in the F 2 the mean 
flower size varies in different plants, also frequently in different flowers 
produced simultaneously on the same plant, and in some cases even the four 
petals of the same flower differ markedly in length. This segregation may 
be either continuous or discontinuous and is clearly non-Mendelian. In 
the F 3 and F 4 generations obtained by inbreeding the same behavior is 
observed, but with a decreasing range of variability. The curve of vari¬ 
ability for a whole F 3 or F 4 population may superficially resemble an ordinary 
fluctuation curve, but it generally shows negative skewness. 

When the curves for the petal length from individual plants are plotted 
they are found to be of many types, some of them showing two, three, four 
or more peaks, others a nearly continuous curve approaching the curve of 
fluctuation, etc. The element of inheritance resulting from the difference 
in size between the flowers of the parent species, the later somatic segrega¬ 
tion on individual plants, and the absence of a normal fluctuation curve for 
most individuals, constitute this a new type variability. Considered as a 

100 


NEW TYPE OF VARIABILITY IN PLANTS 


101 


phenomenon of variability, it differs on the one hand from mutation and on 
the other from fluctuation. Considered as a phenomenon of inheritance, 
it is clearly not Mendelian owing to the irregular character of the segregation 
in the individual as well as between individuals. The condition has arisen 
through failure of adjustment between different size-tendencies inherited 
from the original cross between Oenothera biennis and Oe. rubricalyx. 

Many related species and varieties of plants show conspicuous differences 
in size of flower. If intercrossing occurs in a population of such forms there 
will result a series of types giving a wide range of variaton in flower size. 
This would appear superficially to be due to fluctuation, but would in 
reality be a type of inheritance following crossing. Probably other cases 
of inheritance of petal size when analyzed in the same way as these Oeno¬ 
thera crosses, will be found to show similar non-Mendelian behavior. It is 
possible that the size of repeated parts in general will be found to follow 
the new law in crosses when experiments to determine this point have been 
made. In all such size crosses it will be important to determine the range 
and character of the variation in each individual. 

Since Mendelian behavior appears clearly to be based upon the segrega¬ 
tion of chromosome pairs in the reduction divisions, it appears probable that 
the phenomena of irregular variability and segregation in the individual 
as here described are based upon cytoplasmic differences, since there appears 
to be no definite distributing mechanism involved, and the time of segrega¬ 
tion is not confined to the meiotic divisions. 


THE ANALYSIS OF GENETIC DIFFERENCES THROUGH 
HAPLOID PARTHENOGENESIS 

P. W. WHITING 

The Iowa Child Welfare Research Station , The Slate University of Iowa 

The parasitic wasp, Hadrobracon brevicornis (Wesmael), proves to be 
excellent material for genetic research on account of the ease with which 
it is manipulated, the large number of offspring produced from one female, 
and the brevity of its generation (ten days). The species is strictly 
arrenotokous, virgin females producing males only. Mated females produce 
both males and females, the latter arising in all cases from fertilized eggs. 
In addition to normal females, however, there are produced from fertilized 
eggs, gynandromorphs, mosaic males, and probably other abnormal indi¬ 
viduals incapable of developing to maturity. While mated females may 
give a variety of progeny in peculiar non-Mendelian ratios, ovogenesis 
apparently follows orthodox lines and virgin females produce males from 
reduced eggs in normal gametic ratios. 

Eye-color of the type is jet black. An orange-eyed mutant appeared 
in a fraternity of 253 males from a virgin female. When bred to black, 
orange proved to be completely recessive. Heterozygous females normally 
produce one to one ratio: but a lethal factor has been found showing linkage 
with the orange locus and killing the male pupae which possess it. 

Defects in the fourth branch of the radius vein have been found in the 
progeny of wild females. These abnormalities have proved to be heredi¬ 
tary, but are very variable and much affected by environmental conditions. 
If the wasps are subjected to a rather high temperature during development, 
they are much more likely to show the defect than if temperature is low. 
Well fed large-sized specimens are much more likely to be defective than 
small starved individuals. In general, conditions tending to accelerate 
development produce more defectives, probably on account of a differential 
effect upon wing development as a whole and the development of this 
particular vein. There is also a considerable element of chance in growth, 
as is shown by the asymmetry of the defect, so that if environmental factors 
are constant there is still much variation within homozygous stocks. Hu¬ 
midity appears to be of little importance or none in affecting this char¬ 
acter. Cultures are bred at 30°C. in order to reduce variation due to 
environment. 


102 


ANALYSIS OF GENETIC DIFFERENCES 


103 


Defectives are separated into eight grades, the highest grade (8) denoting 
complete absence of the vein from either wing, the lowest grade (1) denoting 
a very slight break in the vein of one wing. Grade 4 denotes either the 
complete absence of the vein from one wing or its partial reduction in 
both. The frequencies for the different grades may be separated by periods, 
the first position denoting normals. Thus a fraternity of males from a 
virgin female graded,—395.0.2.0.1, there being 395 normal, two of grade 2 
and one of grade 4. 

Certain inbred stocks produce regularly less than 1 per cent defectives. 
These stocks are called low defective. 

Another stock, called type, has produced 2507 normal males and 997 
normal females, besides one defective male (grade 4). This male produced 
normal descendants only and was therefore a somatic variation, not a genetic 
mutation. 

Another stock isolated and bred for a considerable time, produced 93 
per cent defectives. Crosses were made with an orange-eyed stock hav¬ 
ing normal wings and the progeny continued in mass culture. Ten 
females, taken from this culture, produced 149 normal and 154 defective 
males besides a number of females. The mode of the defectives was grade 8. 
The female offspring fell into four classes: (1) Those giving all normals. 
(2) Those giving mostly normal but a few defectives. (3) Those giving 
normal and defective in equal ratio. (4) Those giving chiefly defective. 
A selection for higher defect was started from the fourth group and carried 
through eight generations. Although high grade defective parents were 
selected, there was no modification of the stock, either in ratio of defectives 
or in degree of defect. There were 92 per cent defective in the total series 
and 92 per cent in F 8 . This selected series formed the basis of high defective 
stocks, both black- and orange-eyed. 

An orange-eyed male of grade 8 (od8 d ) from the high defective stock was 
crossed to a female from type stock. Type males and females resulted. 
Some of these daughters were isolated and produced type, defective, orange, 
and orange defective males. One of the last of grade 8 was crossed to a 
type female and daughters again isolated, producing males in ratio as before. 
This artificial alternation of generations may be expressed as follows: 

Type stock 

'nr" 

[—>od8 o' X + 9 = + dV&?9 
-f- 9 = only cf cf,-hi d, o, & od 





104 


P. W. WHITING 


This process was carried to F 2 o, the odd-numbered generations, Fi, F 3 , etc., 
representing the results of sexual crossing, while the even-numbered, F 2 , 
F 4 , etc., are the males resulting from parthenogenetic eggs and show segre¬ 
gation in gametic ratio. The odds, summarized, consist of black males,— 
517.1.0.0.4. and black females,—452.8.16.2.4. The five males with defective 
venation resulted in all probability from fertilized eggs and were sterile 
mosaics. Defectiveness in thirty females probably resulted from chance 
conditions of growth, temperature, etc., which reversed the dominance of 
the allelomorphs so that defective venation appeared instead of normal. 

The evens, segregating generations, consist of black males,— 
899.7.45.40.85.51.112.132.178 or 899 normal and 650 defective; and orange 
males,—921.16.49.42.68.49.99.132.170. or 921 normal and 625 defective. 
This appears to be gametic ratio for two independently segregating factors 
with apparent deficiency of defectives due to somatic overlapping. There 
are in all 1820 normal to 1275 defective. On a one to one basis 272.5 of 
these somatic normals should be genetic defectives. Of the 1547.5 genetic 
defectives, 1275 show the defect or only 82+ per cent instead of the 92 per 
cent as in the selected defective series. Apparently something has been 
taken away that was present in the selected series. It is suggested that this 
may have been the factor for low grade defect, which, when added to the 
high defective factor, gives 92 per cent defective instead of 82 per cent or 
less which may be given by high defective alone. Although the number of 
individuals bred from single females was not large enough to calculate sig¬ 
nificant percentages, there was noticed between F 4 and F 8 a considerable 
decrease in wasps somatically defective in relation to those calculated as 
genetically defective. Both F 2 and F 4 give 87 per cent while the segregating 
generations from F 8 to F 20 inclusive give 78+ per cent. The percentage 
of defectives given by the high defective factor alone may therefore be about 
78 per cent. On the basis of independent segregation of factors for high and 
for low defective, one-fourth of the male offspring of heterozygous females 
resulting from the cross of high selected strain by type should have both 
high and low and 92 per cent of these should be defective, one-fourth should 
have high alone and 78 per cent of these should be defective, one-fourth 
should have low alone and 1 per cent of these should be defective, while the 
remaining quarter with neither defective factor should all be normal. 
The percentage of somatic defectives in the total should therefore be 42.45 
per cent and if one-half of the normals be omitted as has been done in the 
above calculations, defectives make up 85.5 per cent of the remainder. This 
is not significantly different from the 87 per cent found in F 2 and F 4 . F 6 
consisted of numbers too small to be significant, but F 8 gave 78 per cent 
after which there was no further drop. 


ANALYSIS OF GENETIC DIFFERENCES 


105 


An orange male with normal wings was isolated from the first segregating 
generation (F 2 ) of this alternating series and crossed to type females. 
Daughters were isolated and their orange sons again crossed to type. This 
artificial alternation of sexual and parthenogenetic generations was con¬ 
tinued to F 14 . The segregating generations showed no high grade defectives. 
Seven of grade 2 and one of grade 3 appeared in F 2 and one of grade 4 ap¬ 
peared in F 6 after which the factor for low defective was apparently 
eliminated. 

It is obvious from these as well as numerous other studies that, while a 
stock may be uniform genetically, it may show considerable fluctuating 
variability. Whether a genetic character difference from a chosen type 
appears in all of the individuals possessing the genetic potentiality of 
showing it, as is obvious for orange, or in a high percentage but not all, 
as has been practically demonstrated in the case of high defective, or even 
in less than 1 per cent, as may be the case with low defective, the factorial 
basis for the difference may be a simple Mendelian unit. 


MUTATION 


H. J. MULLER 
University of Texas, Austin , Texas 

Beneath the imposing building called “Heredity” there has been a dingy 
basement called “Mutation.” Lately the searchlight of genetic analysis 
has thrown a flood of illumination into many of the dark recesses there, 
revealing some of them as ordinary rooms in no wise different from those 
upstairs, that merely need to have their blinds flung back, while others are 
seen to be subterranean passageways of quite a different type. In other 
words, the term mutation originally included a number of distinct phenom¬ 
ena, which, from a genetic point of view, have nothing in common with 
one another. They were classed together merely because they all involved 
the sudden appearance of a new genetic type. Some have been found to be 
special cases of Mendelian recombination, some to be due to abnormalities 
in the distribution of entire chromosomes, and others to consist in changes in 
the individual genes or hereditary units. It seems incumbent upon us, 
however, in the interests of scientific clarity, to agree to confine our use of 
term mutation to one coherent class of events. The usage most service¬ 
able for our modern purpose would be to limit the meaning of the term to the 
cases of the third type—that is, to real changes in the gene. This would 
also be most in conformity with the spirit of the original usage, for even in 
the earlier days, mutations were conceived of as fundamental changes in 
the hereditary constitution, and there were never intentionally included 
among them cases merely involving redistribution of hereditary units— 
when these cases were recognizable as such. In accordance with these 
considerations, our new definition would be: “mutation is alteration of the 
gene.” And “alteration,” as here used, is of course understood to mean a 
change of a transmissible, or at least of a propagable, sort. 

In thus trimming down the scope of our category of mutation we do not 
deprive it of the material of most fundamental evolutionary significance. 
For all changes due to the redistribution of individual genes or of groups 
of genes, into new combinations, proportions, or quantities, are obviously 
made possible only by the prior changes that make these genes differ from 
each other in the first place. It should in addition be noted that changes 
due merely to differences in the gross proportions of entire groups of genes 

106 



MUTATION 


107 


must be relatively incapable of that delicate adjustment which is required 
for evolutionary adaptation. And as to the question, frequently raised, 
whether all evolution is ultimately due to mutation, this is necessarily 
answered in the affirmative by our definitions of the gene and of mutation, 
which designate the gene as any unit of heredity, and mutation as any 
transmissible change occurring in the gene. The question of the basic 
mechanism of evolution thus becomes transferred to the problem of the 
character, frequency, and mode of occurrence of mutation, taken in this 
precise, yet comprehensive sense. And since eugenics is a special branch of 
evolutionary science it must be equally concerned with this problem. 

In choosing the body of data wherewith to attack these questions of 
mutation, in their new form, it must unfortunately be recognized that the 
results with the evening primrose, Oenothera, although they formed the 
backbone of the earlier mutation theory, can no longer be regarded as 
having a direct bearing on the modern problem, since they cannot be shown 
to be due directly to changes in the genes. Certain of them, such as gigas, 
lata, scintillans, etc., have been proved by Geerts, Lutz, Gates, and others, 
to be due to abnormalities in the apportionment of the chromosomes. Very 
valuable information on the genetics of cases of this sort is now being ob¬ 
tained, especially in the work of Blakeslee, Belling, and Farnham on 
much clearer cases of similar character in the Jimson weed, and, finally, in 
work of Bridges on the fruit fly Drosophila. Most of the other so-called 
mutations in the evening primrose appear to be due to the normal hereditary 
processes of segregation and crossing over, working on a genetic constitution 
of a special type. Evidence for this was obtained in my analysis of the 
analogous case existing in the fly Drosophila, as follows. It had previously 
been shown by de Vries, and further elaborated by Renner, that germ cells 
or individuals of Oenothera bearing certain genes always died, in such a way 
that all the surviving individuals were heterozygous (hybrid) in regard to 
these genes. I later showed, through work on Drosophila, that when such 
a condition (there called “balanced lethal factors”) exists, the situation 
tends to become still further complicated through the presence of other 
heterozygous genes, which are linked to those which cause death. When 
one or a group of these non-lethal genes crosses over (separates) from the 
lethals, as they occasionally do, they may become homozygous, producing 
a visible effect. Thus new types of individuals appear which may be 
ascribed to “mutation, ” whereas they are really due to crossing over. The 
work of Frost on stocks has shown that a precisely analogous situation exists 
in that form also, and G. H. Shull is obtaining direct evidence for the same 
conclusion in the evening primrose itself. In any event, it must be granted 


108 


H. J. MULLER 


that so long as this interpretation cannot be definitely refuted, these varia¬ 
tions cannot be used as examples on which to base our theory of gene change. 
In place, then, of the elaborate system of conclusions which has derived its 
support chiefly from the results in the evening primrose, it will be necessary 
for our present theory of gene change to erect an independent structure, 
built upon an entirely new basis. 

The data upon which the new theory must be built consist of two main 
sorts, which may be called direct and indirect. (1) In the cases giving 
the direct evidence, the occurrence of the gene change can be proved, and 
it is possible to exclude definitely all alternative explanations, such as 
contamination of the material, emergence of previously “latent” factors, 
non-disjunction, etc. So far, the only considerable body of such evidence 
is that gotten in the Drosophila work, where mutations have (in this sense) 
been actually observed in at least 100 loci. Considered collectively, how¬ 
ever, there exist in other organisms enough scattered data to afford ample 
corroborative evidence for the generality of occurrence of mutations like 
those observed in the Drosophila work. In addition several specially 
mutable genes have been found in a number of plants (as well as in Droso¬ 
phila) that are giving highly valuable information along their particular 
lines. And a number of selection experiments that have been performed 
on non-segregating lines of various organisms have also given us direct 
evidence, if not of the frequency, then at least of the infrequency, of muta¬ 
tions. (2) As for the indirect data, these may be gotten by examination of 
Mendelian factor-differences of all kinds, on the assumption that they must 
have arisen through mutation. Although this assumption can be shown to 
be fully justified, these cases cannot provide information concerning the 
manner of origin of the mutants, nor can they furnish a reliable index of the 
frequency of mutations, since the mutant genes may have been subjected 
to an unknown amount of selective* elimination or selective propagation 
before the observations were taken. As for the still more indirect data, 
derived from studies of phylogenetic series and comparisons between 
different species, genera, etc., these occasionally give suggestive results, 
but where crosses cannot be made or where the differences cannot be traced 
down to the individual genes, such facts can seldom lead to trustworthy 
genetic conclusions. 

On these various data, duly weighted, we may found our new mutation 
theory. We know nothing, as yet, about the mechanism of mutation, or 
about the nature of the gene—aside from the fact that nearly all genes 
hitherto studied behave like material particles existing in the chromosomes. 
Nevertheless there is always evidence for a number of empirical principles 


MUTATION 


109 


regarding the changes of the genes, some of which may conveniently be listed 
here in the form of 14 statements. I shall have opportunity merely to 
present these principles, without attempting any adequate explanations of 
how they have been derived from the data. 

1. The first and probably most important principle is that most genes— 
both mutant and “normal”—are exceedingly stable. Some idea of the 
degree of this stability may be obtained from some quantitative studies of 
mutation which Altenburg and I have made in the fruit fly Drosophila. It 
may be calculated from these experiments that a large proportion of the 
genes in Drosophila must have a stability which—at a minimum value—is 
comparable with that of radium atoms. Radium atoms, it may be re¬ 
called, have a so-called “mean life” of about two thousand years. 

2. Certain genes are, however, vastly more mutable than others. Forex- 
ample, a gene causing variegation in corn, studied by Emerson, and another 
in the four-o’clock, studied by Maryatt, ordinarily have a mean life of only 
a few years; and that causing bar eye in Drosophila has a mean life of only 
about 65 years, as is shown by the results of Zeleny. (In expressing these 
results we are here using the physicists’ index of stability, which seem6 most 
appropriate for the present purpose also.) 

3. External agents do not ordinarily increase the mutability sufficiently 
(if at all) to cause an obvious “production” of mutation. 

4. The changes are not exclusively of the character of losses; this is shown 
by the well established occurrence of reverse mutations, in bar-eyed and 
white-eyed Drosophila, in Blakeslee’s dwarf Portulaca, Emerson’s varie¬ 
gated com, and probably in a number of other recorded instances. It is 
known that mutations having an effect similar to that of losses do occur, 
however, and they may be relatively frequent. 

5. The change in a given gene is not in all cases in the same direction, 
and it does not even, in all cases, involve the same characters. The latter 
point is illustrated by a series of mutations which I am investigating in 
Drosophila, which all involve one gene, but which produce, as the case 
may be, either a shortened wing, an eruption on the thorax, a lethal effect, 
or any combination of these three. 

6. The direction of mutation in a given gene is, however, preferential, 
occurring oftener in some directions than in others. This is well illustrated 
in the studies on variegated corn and four-o’clocks, and on the bar eye and 
white eye and other series in Drosophila. 

7. The mutability and preferential direction may themselves become 
changed through mutation, as illustrated by some of the same cases. 

8. The mutations do not ordinarily occur in two or more different genes 
at once. In only two instances in Drosophila have mutations been found in 


110 


H. J. MULLER 


two different, separated 1 genes in the same line of cells of one individual. 
But a recurrent case, apparently of this kind, has recently been described in 
oats, by Nillson-Ehle. 

9. Not only does the mutation usually involve but one kind of gene— 
it usually involves but one gene of that kind in the cell. That is, the 
allelomorphs mutate independently of one another, just as totally different 
genes do. There is evidence for this derived from corn, Portulaca, and 
Drosophila. 

10. Mutations are not limited in their time of occurrence to any particular 
period of the life history. This has been proved in the above mentioned 
studies on mutable plants, in Drosophila, and in other cases. 

11. Genes normal to the species tend to have more dominance than the 
mutant genes arising from them. This is very markedly the case in Droso¬ 
phila, where even the relatively few mutant genes that have been called 
dominant are very incompletely so, and might more justly be called reces¬ 
sive. In other organisms, the same condition of things is strongly sug¬ 
gested, although the direct data on occurrence of mutations is as yet too 
meagre to allow of certainty. 

12. Most mutations are deleterious in their effects. This applies not 
only to the organism as a whole but also to the development of any particular 
part: the delicate mechanisms for producing characters are more likely to 
be upset than strengthened, so that mutations should more often result 
in apparent losses or retrogressions than in “progressive” changes. This is 
both an a priori expectation and a phenomenon generally observed. 

13. Mutations with slight effects are probably more frequent than those 
with more marked effects. This must not be understood as referring to 
the different mutations of each given gene, but it applies in a comparison of 
the mutations occurring in different genes. Thus, there are more than a 
dozen mutations, in different loci, which reduce the size of the wing in 
Drosophila so slightly as to leave it more than half its original length, 
whereas only four reduce it to less than half-length. Mutant genes with 
effects so slight as to be visible only by the aid of specific co-genes seem to 
arise still more frequently. It is reasonable to conclude that the mutations 
with slighter effects would more often take part in evolution, because they 
should usually be less deleterious, and this conclusion is borne out by ob¬ 
servations on the multiplicity with which such factor-differences with rela¬ 
tively slight effects are found in species crosses. 

14. The range of those mutations which are of appropriate magnitude 
to be visible is probably very small, in comparison with the entire “spec- 

1 Contiguous genes may be affected in the rare cases known as “deficiencies,” found by 
Bridges and Mohr. 


MUTATION 


111 


trum” of mutations, so that there are many more lethals than visible muta¬ 
tions, and probably more subliminal than visible. 

The above empirical and semi-empirical principles must be regarded as a 
mere preliminary scaffolding, for the erection of a later, more substantial, 
theory of mutation. Time does not permit me here to discuss which direc¬ 
tions of research, and what methods, seem the most promising for future 
results. Suffice it to say that it is especially important to obtain accurate 
data concerning the effect of various conditions upon the rate of mutation. 
This seems one of the logical routes by which to work towards the artificial 
production of mutation and consequent more perfect control of evolution. 
At the same time such results should also give a further insight into the 
structures of the gene. The way is now open, for the first time, to such 
studies on mutation rate, first through the finding, by Emerson, Baur, 
Maryatt, Zeleny, and Blakeslee, of a number of specially mutable factors in 
different organisms, and second, through certain special genetic methods 
which I have elaborated in Drosophila, for the detection of lethal and other 
mutations there. 

It has now become recognized that advances in theoretical or “pure” 
science eventually carry in their train changes in practice of the most far 
reaching nature—changes which are usually far more radical than those 
caused by progress in the applied science directly concerned. It may there¬ 
fore be asked at this point by eugenists: “Are there any applications 
of the knowledge which has already been gained about mutation in general, 
to eugenics and to the principles which should govern us in guiding human 
reproduction?” I think that one such application is already clearly 
indicated. 

In order to understand the nature of this application it will be necessary 
first to consider the proposition—emphasized by East and Jones in their 
book, “Inbreeding and Crossbreeding”—that the only way for a genetically 
sound stock to be formed is by its going through a course of inbreeding, with 
elimination, by natural or artificial selection, of the undesirable individuals 
that appear in the course of this inbreeding. The truth of this proposition 
depends upon the fact that many recessive genes of undesirable character are 
apt to exist in a population. Since the frequency with which these genes 
are able to produce their characteristic effects, i.e., to “come to light,” 
depends on the closeness of the inbreeding, it is evident that inbreeding will 
be necessary in order to recognize the genes adequately, and hence to elimi¬ 
nate them. 

Our present theory of mutation, however, carries us further than the 
proposition just considered. It shows that these undesirable genes have 
arisen by mutation; in fact, as stated in point 12, the great majority of muta- 


112 


H. J. MULLER 


tions are deleterious, probable even to the degree of being lethal, and it is 
also known, as noted in point 11, that many—probably the great majority— 
are recessive. In other words, our mutation theory shows that probably 
the majority of the mutations that are occurring are giving rise to genes of 
just the type specified in the above discussion. This immediately shows us 
that not only are inbreeding and selection desirable for raising the genetic 
level of a population, but they are absolutely necessary merely in order to 
maintain it at its present standard. For the same process of mutation which 
was responsible for the origination of these undesirable genes in the past 
must be producing them now, and will continue to produce them in the 
future. Therefore, without selection, or without the inbreeding that makes 
effective selection possible, these lethals and other undesirable genes will 
inevitably accumulate, until the germ plasm becomes so riddled through 
with defect that pure lines cannot be obtained, and progress through selec¬ 
tion of desirable recessive traits can never more be effected, since each of 
them will have become tied up with a lethal. To avoid such a complete 
and permanent collapse of the evolutionary process, it is accordingly neces¬ 
sary for man or nature to resort to a periodically repeated, although not 
continuous, series of inbreedings and selections in the case of any biparental 
organism. 

This conclusion is more than a mere speculation, or even a deduction 
from our principles. The reality of this process of mutational deterioration 
has been directly proved, in the case of Drosophila, through experiments 
that I have conducted on lines in which the processes that usually ac¬ 
company inbreeding and selection were prevented: in these lines there was 
found an accumulation of lethal genes so rapid that it would have taken but 
a few decades to have brought about the presence of a lethal gene in prac¬ 
tically every chromosome of every fly. Although the same general thesis 
undoubtedly applies also to mankind we do not yet know the speed of the 
process here. Its speed depends upon the actual frequency of mutations, 
which it will be very important—and extremely difficult—to determine in 
the case of mankind. Meanwhile, no matter what this rate may be, the 
process remains a real one, which must eventually be reckoned with, and 
either grappled in time, and conquered, or else yielded to. 

I have dwelt at length upon this particular application to eugenics, of 
some of the mutation studies. I believe, however, that this is but one ex¬ 
ample of such applications, and that from an increasing knowledge of our 
theoretical science there will inevitably flow an increasingly adequate 
technique for coping with our refractory human material. Meanwhile, the 
crying need is for more of the theoretical knowledge—and for the support of 
pure science, in its investigation of the processes lying at the root of the 
germ plasm. 


REVERSE MUTATIONS 


CHARLES ZELENY 
University of Illinois 

The occurrence of reverse mutation is no longer questioned but cases of it 
are of importance because of their bearing on the general nature of mutations. 
The speaker has worked on the bar-eye allelomorphic series of Drosophila 
which has a number of valuable characteristics: (1) the details of the origin 
of its components are known, (2) the ommatidial counts give a quantitative 
evaluation, (3) the heterozygotes may be recognized and (4) the mutations 
are frequent enough to give an adequate measure of rate. 

About 60 mutants were observed among 150,000 individuals. 

Of the six possible shifts all except full to ultra-bar are on record. 

Reverse mutations are more frequent than the original ones. 

Recent origin is not an explanation of the difference in rate between re¬ 
verse and original mutations. For instance, full derived from bar does not 
mutate more frequently than the original full. 

The large jump from ultra-bar to full is as frequent as the smaller jumps 
from bar to full and ultra-bar to bar. 

Direction of selection has no effect upon the rate or direction of mutations 
as observed in 42 generations of upward and of downward selection for eye- 
facet number. 

Mutation outside of the bar locus may affect eye-facet number but there 
is no evidence that the presence or absence of such accessory mutations 
affects the rate or direction of mutation at the bar locus. 

Direction of origin has no effect upon the direction or rate of mutation. 
For instance bar derived by upward mutation from ultra-bar differs in no 
respect from bar derived by downward mutation from full. 

There is no evidence of periodicity in the mutations. 

Mutations occur in the germ plasms of both males and females. This is 
indicated by the preponderance of females among the mutants, bar being a 
sex-linked character. 

The mutations are not confined to a single period in the germ-cell history. 
Some of them at least come before the end of the gonial divisions because 
five of the forty-four cases valid for this purpose show mutant individuals 
appearing together. 


114 


CHARLES ZELENY 


The frequency of mutation at the bar locus is greater than in all observed 
accessory factors combined. 

The different components of the bar series are definite entities comparable 
to definite chemical compounds or physical states. They do not grade into 
one another and their origin is not a factor in their behavior. 

So far no environmental factor influencing rate of mutation in bar-eye 
has been discovered. There is no difference between stocks kept at 20° to 
22°C. and those kept at 27°. 

These results have an obvious bearing upon certain theories of evolution 
but there is not time to discuss them on the present occasion. 


DARWINIAN EVOLUTION BY MUTATIONS 
R. A. FISHER 

Rothemsied Experimental Station, Harpenden, England 

During the second half of the nineteenth century, the Darwinian theory of 
the gradual evolution of all living forms by the agency of Natural Selection, 
slowly won its way to acceptance, first by men of science, especially breeders 
and geologists, and finally by the whole body of educated opinion. It is 
not here the place to survey to the full extent of this revolution in human 
thought; it is enough that to thoughtful minds it dominated the outlook 
upon the history and destiny of the human race, and gave birth at the hands 
of Francis Gal ton to the Science, or rather the philosophy, of Eugenics. 
But, during the present century Darwin’s views have been exposed to criti¬ 
cism in detail and in gross, from the two classes of students, the breeders and 
the palaeontologists by whom it was first most readily received. It is 
the purpose of the present note to examine very briefly the causes of this 
change of attitude, and to clear away certain misunderstandings, which 
spring chiefly from changes in the use of words, which have taken place 
during the past half century. 

In the first place the discovery of Mendelism has made us familiar with the 
fact that obvious and easily distinguished differences in animals and plants 
are sometimes due to a single heritable factor, and some Mendelians have 
in consequence taken offence at the gradual and cumulative character which 
Darwin assigned to evolution. On the other hand certain palaeontologists 
to whom the gradual and progressive character of the evolution of fossil 
remains is becoming more and more evident, feel that we have here some¬ 
thing which the geneticist cannot explain, and consequently fall back upon 
Lamarck’s suggestion of the inheritance of acquired characters, or upon 
the mystic word “orthogenesis.” Others again, impressed by the genetic 
constancy in pure line breeding, have somewhat rashly insisted that genuine 
mutations never occur. These different views, though proper to put 
forward for discussion among men of science, exert a bewildering effect upon 
the general public, who tend to lose their belief that science has anything to 
teach them about the history and the destiny of their race. 

Of the facts unknown to Darwin and his contemporaries we have to take 
two into consideration. In the first place Mendelism shows not only that 

115 


116 


R. A. FISHER 


obvious and easily distinguished differences may be due to single factors, 
but that the ordinary differences between parent and offspring, or between 
children of the same parentage, may be, and probably are for the most part, 
due to the segregation of Mendelian genes, and not, as Darwin seems to 
have thought, largely to new and arbitrary mutations of a heritable nature. 
What the older evolutionists took to be for the most part new heritable 
differences, the Mendelian interprets as, for the most part, old heritable 
differences, newly arranged according to the Mendelian system. In the 
second place pure line experiments have shown that in genetically pure 
strains, the appearance of entirely new genes is of relatively rare occurrence. 
It is not usually understood that these two new facts are logically con¬ 
nected: for, once the Mendelian view is accepted that genes do not blend, 
but segregate intact, it is clear that if in every generation new genes are 
introduced, the variability of the species will increase without limit. If 
the genes of the parents were to blend in the offspring, continual new muta¬ 
tions would be necessary to maintain the variability; but as under the Men¬ 
delian system of segregation there is no tendency for the variability to 
diminish, except in so far as by the gradual action of selection certain genes 
tend to disappear, we must not assume that in a state of nature, where the 
variability is approximately constant, new Mendelian alterations are in¬ 
troduced into the currency of the stock more rapidly than this gradual 
elimination takes place. 

The fundamental facts upon which Darwin grounded his theory are more 
firmly established than ever: the universal tendency of animals and plants 
to breed up to the limits of subsistence has never been seriously questioned; 
the existence in wild and domesticated races of heritable differences has been 
consistently verified; the incidence of natural selection and the actual modi¬ 
fication of types has been proved by many careful investigations. Never¬ 
theless, owing to the changes which have taken place in the use of words, 
many would feel almost as though they were out of date if they styled them¬ 
selves Darwinians. This change in terminology is principally due to the 
far reaching effects of the factorial system on our ideas of the constitution 
of living things. There is no need here to discuss the proper use of modern 
terms; for the purposes of the present note it will be sufficient to say that we 
shall speak of a species as differing in any Mendelian factor, when two 
or more allelomorphs of that factor are to be found in individuals of the 
species; that every individual of the species must belong, in respect of this 
factor, to one or other of the homozygous and heterozygous types formed by 
combining like or unlike allelomorphs; that the word gene will be used for 
the material basis of any allelomorph, and the word locus for the material 


DARWINIAN EVOLUTION BY MUTATIONS 


117 


basis of a factor: so that we may speak of one gene supplanting another in 
the same locus as one allelomorph replaces another of the same factor. On 
the factorial system, then, an individual is specified if, for every factor 
concerned, we assign it to one or other of the homozygous and heterozygous 
types: in the simplest case, when the factor is dimorphic, there being only 
two homozygous and one heterozygous types. Any organ or trait of the 
individual will usually be influenced by many factors, so that the selection 
of any one trait will influence the proportions of the allelomorphs of all 
the factors which affect that trait. Since in nature many traits are, or more 
correctly the whole complex of traits is, subject to selection, natural selec¬ 
tion within any species will necessarily be gradually increasing the propor¬ 
tions of some allelomorphs, and diminishing those of their alternatives, so 
that a gradual progress of the whole specific group must take place on the 
whole in the direction of improved adaptation to those needs which domi¬ 
nate selection. 

The action of selection upon such a species will be exactly what the Dar¬ 
winian would anticipate, with the theoretical reservation that in the absence 
of any mutations, progress must sooner or later stop; for the number of 
possible types formed by combining all the possible allelomorphs of all the 
factors present, though inconceivably great, is still finite. Of these types 
one is presumably the best adapted to the selecting environment, and when 
that type is attained improvement ceases. Of course the best possible com¬ 
bination of factors may not, when selection commences, exist in a popula¬ 
tion of many millions; thousands of generations of severe selection may be 
needed to bring it into existence, and establish it as the dominant type; 
it is true also that in the absence of new genes, the average value of any trait, 
such as human stature, might be changed to a value far outside the existing 
range of variation, merely by selection. Still it must be admitted that in 
the absence of mutation the variability of the selected species would be 
progressively diminished, and will finally vanish, so bringing evolutionary 
progress to an end. 

But modern work, especially that of American workers on Drosophila, 
shows conclusively that mutations though infrequent do in fact occur; 
and it is worth while to observe exactly what bearing this fact has upon the 
Darwinian theory of evolution. 

If we suppose then that a mutation has occurred, and an entirely new 
gene is present in a single individual of a population consisting of some thou¬ 
sands of millions, the history of its survival maybe broadly divided into two 
periods. In the first period its survival or extinction is due mainly to 
chance, in the second period mainly to the general advantage or disadvan- 


118 


R. A. FISHER 


tage in the struggle for existence which the new allelomorph confers, on the 
average and in combination with the existing currency of genetic types, as 
compared with the alternative allelomorph, which it displaces. 

In consideration of the first stage we may suppose that the chance of any 
gene of one individual appearing in 0, 1, 2, 3 . . . . individuals in the sec¬ 
ond generation, to be po, pi, P2, P3, . . . • etc., such that 

Po + Pl + P 2 4-.=1 

These fractions will depend on the stage in the life history of the individual 
which we pick out for consideration: for the adult reproductive stage of 
many plants and animals, the series will be very similar to the Poisson 
series 


If we construct a function 

f (x) = po -f Pi x + p 2 x 2 . 

then the chance of any one gene being represented in the second filial gen¬ 
eration by 0, 1, 2, 3 ... . individuals will be found by substituting 
f(f(x)) for f(x). 

This method enables us to compute the chance that the gene will not have 
become extinct in any number of generations; assuming the Poisson series, 

2 

the chance of survival for n generations is nearly while if it do survive 

the average number of individuals affected will be |n, Thus roughly one 
mutation in 50 will survive 100 generations, and if so, it will on the average 
be represented in 50 individuals. These results are worked out for a popu¬ 
lation stationary in number. 

Very disadvantageous genes, such as dominant lethals, will of course be 
cut off at once, but for those which are only of moderate advantage or dis¬ 
advantage, the above may be taken to represent the first stage in survival, 
which is principally governed by chance. The second stage commences 
with the new gene established in a fairly large group of individuals, of vary¬ 
ing genetic constitution, so that a fair number of new genetic combinations 
are being tested simultaneously. If those individuals which contain the 
new gene are found on the average of the chances of life, and on the average 
of the genetic natures with which it is combined, to be at a disadvantage in 
the struggle for existence, then the number of the mutant form will gradu¬ 
ally diminish, with large fluctuations due to chance; in this way the disad- 





DARWINIAN EVOLUTION BY MUTATIONS 


119 


vantageous gene will always be kept sufficiently rare to be in danger of 
extinction, and though the number bearing it may repeatedly be reduced 
very low without actual extinction, yet sooner or later fortune will fail it, 
and it will disappear from the race. On the other hand the gene which is 
found to confer a slight average benefit on the individuals bearing it, 
will tend to increase in numbers, somewhat more rapidly than its less 
favourable allelomorph. Even if the average advantage be only of the order 
of 1 per cent in a generation, it will gradually spread through the popula¬ 
tion. At first the fluctuations from year to year will be large; so that 
when 100 individuals are affected the average increase will be one in a gen¬ 
eration with a standard deviation of 10. But when the number of affected 
individuals is larger the increase takes place with greater and greater regu¬ 
larity; for example a population of 1,000,000 will have an average in¬ 
crease of 10,000 with a standard deviation of 1000. At this point the 
spread of the favoured gene takes place with calculable regularity. Finally 
when nearly the whole population is affected its less favourable allelomorph 
becomes sufficiently rare for its survival or extinction to be at the caprice of 
fortune. 

If we have rightly described the manner in which a new mutation is 
incorporated into the general stock of the hereditary qualities of a species, 
and the manner in which the variability of the species is maintained in 
spite of the occasional extinction of genes by selection, we are in a position 
to see how great an advantage it is to a species to have adopted methods of 
sexual reproduction on the Mendelian system. For mutation is necessarily 
a leap in the dark: the chances of failure are far greater than those of suc¬ 
cess, especially when the effect of the mutation is large. Hence there is a 
great deal to be gained if it be possible to maintain the variability of the 
species, with a minimum of mutations,—that is to say with the greatest 
stability of the reproductive processes. Now in a population differing in 
a great many Mendelian factors as all sexual populations are found in 
nature to do, a single mutation may enable thousands of new genetic com¬ 
binations to be tested, and if any of these should happen to be very advan¬ 
tageous, it will by selection become the predominant type. It cannot be 
denied that many groups of animals and plants appear to carry on success¬ 
fully by asexual methods of reproduction, but it is impossible to believe that 
these could adapt themselves so rapidly to slow changes of environment as is 
possible with sexually reproductive organisms. 


THE FREQUENCY OF MUTATION AND THE INCIDENCE OF 
HEREDITARY TRAITS IN MAN 


C. H. DANFORTH 

Washington University School of Medicine, St. Louis , Missouri 

In man, as in lower forms, pairs of somatic traits which show the Men- 
dclian type of heredity have presumably come into existence as a result of 
mutation, one of the allelomorphic genes having been derived from the other, 
or both genes having arisen from the same source. The current conception 
of the gene as a discrete and more or less specific determining unit which is 
passed along through the germplasm of successive generations unchanged 
except for occasional mutations invests the question of the nature and fre¬ 
quency of such mutations with a fundamental importance. But while it 
has been possible for a number of investigators to secure rather precise 
evidence as to the frequency of several mutations in experimentally con¬ 
trolled material, data from human sources have thrown very little light on 
mutations in man. There are, however, some considerations which, if they 
do not lead to precise information, do at least serve to indicate the upper or 
lower limits for the frequency of certain mutations in the human germplasm. 
It is the purpose of this paper to present a brief discussion of some of these 
points. 

The theory of the gene as generally understood implies the persistence of 
the same elements through successive generations, and the mathematical 
theory of probability leads to the expectation that these elements will persist 
in the same relative proportions so long as no differential factors come into play. 1 

The two factors which might change the proportions are Selection and 
mutation. If the ratio between two alternative genes is known and the 
selection factor can be evaluated, a means is afforded for estimating the rate 
of mutation. This has been attempted with two classes of hereditary traits, 
in one of which the selection factor is presumably effective, but adversely, 
while in the other it is apparently negligible. By the term selection must 
be understood in this connection any influence whatsoever which may act on 

1 This matter has been discussed very fully by Jennings (Genetics 1: 53-89 and Genetics 
2:97-154), Wentworth and Remick (Genetics 1: 608-616) and Wright (Genetics 6: 111— 
123, 167-179). 


120 


FREQUENCY OF MUTATION IN MAN 


121 


either germplasm or soma in such a way as to affect differentially the rate of 
reproduction of two alternative types. Among such possible factors might 
be, for example, differences in the “survival value” of the two types, selec¬ 
tive mating, linkage with favorable or unfavorable traits, or any other 
agency which might affect the two classes of individuals differently. 

As representatives of traits which might be subject to selection that type 
of polydactyly which is characterized by a (frequently rudimentary) digit 
on the ulnar side of the hand or the lateral side of the foot, and the form of 
syndactyly in which the third and fourth digits of the hand or the second 
and third of the foot are united, were selected for consideration. These 
two traits are distinct and behave in heredity as dominants. They are 
frequently simulated by pathological conditions which are not hereditary 
or represent by-products of some other hereditary condition. There is 
evidence of one or more rather rare genes in the human germplasm which are 
capable of suppressing polydactyly in an individual where it might be 
expected to appear. Bonnevie 2 seems to have found such an instance. 
But a study of family histories makes it quite clear that the presence of this 
trait is due primarily to a single gene which is not dependent upon the coin¬ 
cident action of other genes (except those for which the species is homozy¬ 
gous) and which is only rarely entirely suppressed by other hereditary 
factors. A similar analysis of the data on syndactyly leads to similar 
conclusions in regard to the gene for that trait. 

The present frequency of the genes for these traits can be determined only 
from the incidence of the traits themselves and the evidence on this point 
is rather unsatisfactory. According to records in the office of the Surgeon 
Genera lof the United States Army there were, among a certain ten thou¬ 
sand soldiers demobilized at Camp Dix, ten who showed syndactyly. In all 
of these cases the condition was confined to the feet. Those individuals in 
whom the syndactyly occurred also in the hands had doubtless been refused 
admission to the army. This would seem to indicate a higher incidence in 
the population as a whole, but the probable error is rather large and such 
data as it has been possible to collect from other sources indicates that the 
incidence of this trait can not be much greater than one in a thousand. A 
comparison of the frequency of cases recorded from the clinic indicates that 
the incidence of polydactyly is slightly, but only slightly, higher. 

Since the traits are dominant and marriage between two similarly affected 
individuals is extremely rare, most cases indicate heterozygosis and the 
incidence of the genes is approximately one-half the incidence of their 

* Bonnevie, Kristine. Polydaktyli i norske bygdeslegter. Norsk Mag. f. Lagen., 
1919, pp. 1-32 (known to the writer only through reviews). 


122 


C. H. DANFORTH 


respective traits. With matings strictly at random the precise ratio between 
a gene and its allelomorph could be determined if the exact incidence of the 
trait were known, for it has been frequently pointed out that if under such 
conditions the ratio between two allelomorphic genes is as a: b the proportion 
of the different classes of individuals is as a 2 homozygous for one condition, 
to 2ab heterozygous, to b 2 homozygous for the alternative condition. In 
the present instance (a 2 + 2ab): (a 2 + 2ab + b 2 ):: 1:1000, from which the 
values of a and b may easily be deduced. Determined either way it appears 
that in the germplasm of the whole population there is about one gene for 
syndactyly to every two thousand genes for the allelomorphic normal 
condition. 

A matter of especial interest at this point is the fact that both polydactyly 
and syndactyly occur in widely different racial stocks and have also been re¬ 
ported in different species of anthropoid apes, not to mention still lower 
forms. In view of the great diversity of conditions under which present 
races and species exist it is difficult to believe that the appearance of these 
traits is anything new and the conclusion is justified that they have recurred 
in different lines during immense periods of time, perhaps antedating the 
origin of many existing species. If either of these traits had any positive 
value in increasing the ultimate fertility of its possessors one might expect 
that it would long since have become common to the whole species. Before 
proceeding beyond this point, however, it is desirable to inquire just what 
should be expected of a trait which has a positive selection value. 

For the purpose of such an analysis it may be assumed that at a given time 
the ratio between the number of genes representing a dominant trait and 
the number representing the normal allelomorphic condition is as a to b. 
Then if individuals possessing the trait were to mate always with normal in¬ 
dividuals, the number showing the trait would be in the proportion of a 


heterozygotes to 


b—a . 

—normal recessives. 


The subsequent matings will 


b —a 

, . 2 b — 3a 

be a between heterozygous and normal individuals and--— = —~— 

2 4 

between normal individuals. If m be taken to represent the average 
number of offspring for each mating, the first group will produce hetero¬ 


zygous and ^ homozygous offspring, while the second group will produce 
m (b-3a) 

- , all homozygous and normal. The ratio between the number 





FREQUENCY OF MUTATION IN MAN 


123 


of offspring that carry the trait and those that do not is ~ •’ — -f 
ma 

~n~ ~ 2 ma:m (b—3a) + 2 ma = 2a: (b—a). In the absence of selection 


this ratio would represent the relative proportion of heterozygous to 
normal individuals in the second and succeeding generation, but if the 
traits have a different survival value, then a certain proportion, s, of 2a and 
a different proportion, t, of (b—a) will survive and the ratio of children 
born in the second generation will be altered to 2sa: t(b—a). If hetero¬ 
zygous individuals were always to mate only with normals this relation 

2 s n a 

would assume the general form— b _ a y where n is the number of generations 


considered. When the mutation confers any advantage, s is greater than t 
and the formula represents the minimum possible rate of increase for the 
incidence of the trait, since it provides that through heterozygosis a maximum 
number of recessive normal genes shall profit by the favorable selection 
occasioned by the effect of the dominant genes with which they are asso¬ 
ciated. This formula applies up to the time when heterozygous individuals 
become sufficiently numerous to occasionally mate with each other. If 
individuals showing the trait were always to mate with each other a condi¬ 
tion approaching homozygsis would obtain and the ratio of mutant to 

as n bt n as 11 

normal individuals would be as — : ° r simply —. This formula 


represents the maximum possible rate of increase. With random mating 
the true value of the ratio, of course, lies between these two extremes. 
But when, as is usually the case, a is very small and b is very large the value 
of the two fractions is almost identical until the ratio of numerator to 
denominator reaches about 1:10. It will be apparent, from the nature 

2 s n a 

of the matings it presupposes, that the value of t n^_ a j can never 


exceed 

Two steps serve to reduce these formulae to more practical form. First 
by converting s:t into such form that t is equal to 1, t D may be eliminated 
from the denominator. Second, what is really desired is to know how 
many generations will be required to raise, by selection, the incidence from 
a a' 

— to The second formula developed above represents the relative inci¬ 


dence both of mutant to normal individuals and of mutant to normal genes, 
the first formula represents the ratio between heterozygous and normal 
individuals. Converting the first formula into a formula for the ratio of 





124 


C. H. DANFORTH 


mutant to normal genes we have 2 as n mutant genes to 2 (b—a) + 2 as" 


as 1 


we 


normal genes = as": (b — a) + as". Putting ^ _ a ) -f- as" “ b” 
have b' as" = a' (b — a) -f a' as" or b' as" — a' as" = a' (b — a), whence 


a' (b — a) 


a'b 


s - = ——-- In the second formula s" = 77“ • The values of all 

a (b' — a') b a 

the terms in these equations are known, or assumed, except the value for 

n (number of generations) and this value is represented by 

, r [V (b-a)-l 

n _ log - J, 

log. s. 


in the one case and 




in the other case. 


In other words, if the ratio of two genes is as a:b and the one gene confers 
upon its possessor qualities such as to give an advantage over the alterna¬ 
tive condition that may be represented as s:l, the number of generations 

a a' 

required to raise the original incidence from — to jy will be somewhere 

between the values represented by these two formulae. As stated above 
a' 

so long as j “7 is not greater than T V, the difference between the two values 

amounts to almost nothing. Fortunately in all ordinary circumstances 
this requirement is met and either formula alone is serviceable. 

It is not known what the incidence of polydactyly or syndactyly may 
have been in any past epoch but the formulae developed above show that 
had any dominant trait had an incidence of one in a million and had it con¬ 
ferred in the long run an advantage one one-hundredth greater than the 
advantage conferred by the recessive normal condition it would have taken 
only about 925 generations for the trait to acquire a frequency ten times as 
great as the present frequency of syndactyly. As indicated above there is 
evidence that both syndactyly and polydactyly have been in existence for 
a period many times greater than that indicated by this number of genera¬ 
tions, from which it may be inferred that neither of these traits has had 
any positive selective value. The possession of these traits has been either 
a matter of indifference or, in the long run, a positive detriment to the lines 
in which they occurred. In the former case the present incidence of the 
genes for these traits represents the accumulated mutations which have 
occurred during the past history of the race and there would seem to be no 
escape from the conclusion that since earliest times one line of genes in 







FREQUENCY OF MUTATION IN MAN 


125 


approximately every two thousand has undergone mutation from the normal 
to the syndactyly producing form. The number of lines that have mutated 
to the polydactyly producing form is apparently slightly greater. Since 
the time when the first mutation occurred is unknown this throws no light 
on the rate of mutation beyond indicating that it must have been extremely 
low. 

If the trait is actually unfavorable in its net effect, the rate of mutation 
must be such as to bring the incidence to its present value and to balance 
the adverse effect of selection. In a population numerically about station¬ 
ary the average number of generations through which a dominant trait 
persists among any descendants of a person showing the trait gives a measure 
of the selective value of the trait. Letting n represent the average number 

a 

of generations through which a trait persists in a family and — the inci¬ 
dence of the trait in the population at large it follows that the frequency of 

a 

mutations necessary to maintain that incidence will be “. There is a 

rather common belief, and possibly some inconclusive evidence, that both 
polydactyly and syndactyly represent, or are in some way associated with 
unfavorable factors, but an examination of a considerable body of data 
makes it seem highly improbable that the average duration of these traits 
within the families where they occur is less than three generations even when 
correction is made for the rapid increase in our population. With n = 3 
and a:b = 1:2000 the frequency of mutation is indicated on an average of 
one gene in six thousand. Mutations from the normal to the syndacty] 
condition can hardly be more frequent than this unless it be that the con¬ 
dition is actually increasing or that when the mutation does^ appear it per¬ 
sists, on the average, for less than three generations. If, as is quite possible, 
syndactyly has an average duration of more than three generations the 
frequency of mutation must be correspondingly less. The data for poly¬ 
dactyly, as indicated above, are essentially similar. 

It may be recalled that there is a considerable number of dominant 
traits which are probably slightly unfavorable and which have an incidence 
not greatly different from that of syndactyly. The incidence of these 
traits is no doubt maintained in part by recurring mutations. .The fre¬ 
quency of such mutations could be estimated if the average number of 
generations through which they persist were known, but it is very doubtful 
if the maximum frequency is often greater than 1:6000. In some of these 
there is evidence that the average duration is for only a very few generations, 
and in such cases the maximum frequences of mutations can not be much 
below this value. 


126 


C. H. DANFORTH 


The second class of traits mentioned at the beginning, those in which the 
selection factor is apparently negligible, admits of a slightly different treat¬ 
ment. The traits selected from this group as types are (1) suppression of 
the palmaris longus muscle, apparently a dominant trait, 3 and (2) complete 
absence of hair from the middle of segments of the digits, a recessive trait. 4 * 
Both of these traits occur in various races of man and in occasional anthro¬ 
poid apes. A similar analysis to that employed in the case of syndactyly 
shows that neither of these traits has any appreciable value in increasing 
the differential productivity of its possessors. On the other hand there is 
not the slightest indication that they are in any way disadvantageous, and, 
being so obscure that they can be detected only by careful examination, the 
possibility of sexual selection is also eliminated. 

For purposes of analysis the data on the muscle furnish much the better 
material. Fortunately absence of this muscle has been studied extensively 
in the anatomical laboratories of various countries with the result that 
extensive and reliable data are available. The manifestation of the trait 
is somewhat influenced by sex, being less affected in females, which conse¬ 
quently furnish better material for genetic study. In this sex the trait is 
manifest (i.e., the muscle is absent on one or both sides), according to pub¬ 
lished reports, in 14.7 per cent of bodies dissected at Petrograd and in 21.9 
per cent of bodies dissected at Paris. In Alsace the percentage is about the 
same as in Paris. If the trait is dominant and the reported frequencies are 
approximately correct, the incidence of the gene for suppression of the mus¬ 
cle must be about 10:115 in the population from which the Russian dis¬ 
section material was drawn, and about 16:115 in the population from which 
French material came. In the American negroes the incidence of the gene 
is not over 5 in 115. 

Since there is no positive selective value associated with the trait the 
difference in the incidences for these three groups must be attributed to 
differences in the number of mutations that have occurred. If the existing 
races of Europe were differentiated from a common Asiatic stock toward 
the close of the early Paleolithic period, and other races of men separated 
somewhat earlier in this period, 6 the present European races are separated 
from their common ancestor by some 50,000 years and from the common 
ancestor of all races by perhaps three times that period. Allowing an 
average of four generations to the century this represents respectively about 
2000 and 6000 generations. Attributing the different incidence in the 

3 Thompson, Batts and Danforth: Amer. Jour. Phys. Anthropol. 4: 205-218, 1921. 

4 Danforth: Amer. Jour. Phys. Anthropol., 4: 189-204, 1921. 

6 Osborn: Men of the Old Stone Age. 


FREQUENCY OF MUTATION IN MAN 


127 


different races to differences in the number of mutations that have occurred 
and assuming that the race with the smaller incidence represents approxi¬ 
mately the incidence of the parental stock the difference between two groups 
is an index of the minimum number of mutations that has occurred since 
the two stocks separated. 

Comparing the incidence of the gene in the negro and the Russian germ- 
plasm it appears that for every 115 normal genes there are about 5 more 
mutant genes in the latter than in the former. If 6000 generations repre¬ 
sents anywhere nearly the time since these two groups, along with others, 
diverged from a common ancestor, these additional mutant genes must have 
been acquired in this length of time. If, as is probably the case, the negro 
has been producing occasional mutations during this period, the Russian 
germplasm must have produced slightly more than the number indicated. 
These figures indicate that in the course of 6000 generations at least 5 in 
every 115 gene lines which have persisted, underwent mutation in this one 
direction. Since there is no evidence of selection, this may be presumed 
to be the rate of mutation in this particular germplasm, from which it 
follows that there has been, on an average, one mutation in every generation 
for each 138,000 genes. If, as some might be inclined to think, the two 
germplasms have passed through more than 6000 generations since they 
separated, the rate of mutation has been less than this. 

The white races represented in Europe are more closely related and the 
approximate time of their divergence may be estimated with greater assur¬ 
ance. It is doubtful if these races have during the past 50,000 years re¬ 
produced at an average rate much in excess of four generations to the cen¬ 
tury, in which case 2000 generations may be assumed to be close to the 
maximum time since the separation of the west Russian from the French 
and west German stocks. At the present time there is a difference between 
these stocks of 6 mutant genes to every 115 normal genes. Correcting for 
the rate of mutation in the Russian germplasm as determined above there 
has been in the more highly mutating “French” stock during this period 22 
mutations for every 345 lines that have contributed to the present popula¬ 
tion. This gives an average for each generation of one mutation to every 
31,364 genes. This represents the minimum rate, but there is no reason 
apparent for thinking that the mutations could have been much more fre¬ 
quent than this. In our own population where the incidence of the trait is 
slightly higher than in France there must be with our present birth rate, an 
average of one of two cases each year in which the trait appears de novo. 

In the case of complete absence of hair from the middle segments of the 
digits there are much less extensive data. In white Americans in general 


128 


C. H. DANFORTH 


the ratio of the gene for this trait to that for its normal allelomorph is about 
1:1, while the American negro is almost homozygous for it. These facts 
are not sufficient to warrant an attempt to compute the rate of mutation, 
but it may be pointed out that in this case, unlike the preceding, the rate has 
been much higher in the negro than in the white race. 

In conclusion: It has been shown that when certain conditions are met a 
basis is afforded for estimating the approximate rate of mutations in man. 
Some mutations occur repeatedly, but the frequency of such occurrences 
differs in different racial stocks. Moreover a race which shows a relatively 
low incidence for one type of mutation may show a relatively high incidence 
for another type. These facts indicate a parallelism, but not an identity, 
in the general tendencies of different human germplasms. 


A SIMPLE EXPLANATION OF THE HEREDITARY 
MECHANISM 


LOUIS LEGRAND 
Tunis , Africa 

For half a century most biologists have tried to visualize heredity by 
the means of special representative particles which, hypothetically, were 
endowed with all the qualities needed by their theory. It is possible, 
however, with a minimum of creations of the mind, to obtain a satisfactory 
idea of the hereditary mechanism, by the use of some very simple construc¬ 
tions, without any need of special particles and without leaving the ground 
of observed facts. 

It is first necessary to admit plainly the distinction between the fixed 
and immutable characters and the not fixed and changeable ones, and this 
distinction remains true for the two realms of Nature. 

From the characters which can define individuals and which are con¬ 
stantly transmitted, we must separate the characters which are never seen 
to change in animals of the same species: For example four limbs and hair 
for mammals; retractile nails for a carnivore; a single occipital condyle 
and feathers for a bird. Such characters are the basis of our classifications; 
they are subject to the laws of correlation and are as much immutable as 
they are interesting. Some most important parts of the body of an animal, 
teeth, for example are of this type, which is very often used in the classi¬ 
fications. 

Fixed characters are at least specific characters, and even still more 
comprehensive, being also characters of genus, of order, of class, of branch. 

Other characters, which vary from an animal to another of the same 
species, and from one generation to the following one, ought to be called 
“unfixed characters,” and are to be ranged as follows: 

1. Sex and secondary sexual characters. 

2. Characters of race and variety: for example in the human species, 
the color of the skin, of the hair, of the iris, the pigmentation of epidermal 
cells, the form of hair on section, the cephalic, nasal and maxillary indexes, 
constituting the facial physiognomony of each race; the average stature and 
span; psychic aptitudes of each race; all these characters are always trans¬ 
mitted when parents are of the same race. 

129 


130 


LOUIS LEGRAND 


In the third group, linked with the preceding by insensible shades, we 
find some superficial, sporadic and less important characters, which are not 
regularly transmitted; they stand in the neighborhood of teratology and 
pathology, such as hemophilia, baldness, albinism, familial deafness, dalton¬ 
ism, hereditary optic neuritis; some diathesis compatible with fecundity; 
the coarseness or thinness of face, of hand or foot; musical or mathematical 
aptitudes,— all these are atavic or ancestral characters. 

The last group also includes the quite indefinite variety of characters 
suitable to immediate parents; their transmission sometimes is very precise, 
sometimes not recognizable; their type is like naevi, like some very slight 
details of body, of sensorial apparatus, of skin; like the quality of arterial, 
muscular, glandular tissues; the most part of diathesis and all the scale of 
tempers; some psychoses and neuroses; longevity; some qualities and 
sensorial deficiencies, such as short-sightedness. Such heredity can be found 
to hold even to histological details, as for example the familiar resemblance 
of palmar veinous arcades, and chiefly these dispositions of the papillae 
of the finger’s pulp, so often used for anthropometric test. 

Fixed characters are evidently hugely more important than unfixed ones. 
It is surely an insignificant thing which causes a horse to become white or 
black, swift or bulky, fit for race or for carriage, it is, on the other side, 
something important which causes the same horse to become a vertebrate, 
a quadrupede, an herbivorous, a solipede. 

If such a discrimination between fixed and unfixed characters is admitted, 
we have now to confront it with the constitution and histologic description 
of genital cells. In other words, it is necessary that some concrete and dis¬ 
tinct elements, corresponding to each genus of characters, should be found in 
that plasmatic mass which is*tiescribed at the beginning of each life, that is 
to say the fertilized egg. 

Let us then consider a fertilized egg, already separated from the mother, 
in order to abstract all maternal plasmatic supply: such an egg, by defini¬ 
tion, if kept in a convenient place, will be fit for segmentation, that is to 
say to develop these fixed and unfixed characters according to their recip¬ 
rocal importance and to synthesize them into the well known specific forms. 

The schematized fertilized egg, in whatsoever living type we consider, 
appears like a globular mass of cytoplasm of maternal origin, including two 
half nuclei, one of maternal, the other of paternal origin. As the cytoplasmic 
mass is greater than and surrounding these nuclei, we are logically led to 
believe that cytoplasm is the material support of fixed characters, which 
characters are common to both sexes and also predominant in the features 
of every living being. By exclusion, these two half nuclei, of comparatively 


EXPLANATION OF HEREDITARY MECHANISM 


131 


such reduced size, will correspond to unfixed characters, which are, or may 
be different in each of the parents. 

So it is no question here of using some strange or scarce histological 
peculiarity; we only rely upon what is universal and known for a long time, 
namely, these three distinct elements at the origin of each ontogenesis. 
One is common to both sexes and is this large mass of plasmas (or characters) 
fixed together during all the life of species and of individuals. This is 
temporarily fused with two small masses of plasmas (or characters), un¬ 
fixed during the life of species, but fixed henceforth during the individual 
life. It is to be believed that such a disposition is the result of some physio¬ 
logical division of labor between both sexes, as ancient in phylogeny as is 
the appearance of the Metazoa. 

We must now try to analyze further this schematic explanation of the egg, 
and realize the mechanical relations by which unfixed plasmas ought to be 
ruled. The same gradation which has been admitted in the way of fixation 
of characters can be found again in the respective spaces occupied by their 
representative plasmas, in the egg ready for segmentation. In other words, 
unfixed plasmas will then be nearer to the position of the definitely fixed 
ones (that is to say to the surrounding cytoplasm) since they will repre¬ 
sent some the more anciently and some the more recently fixed characters. 
Consequently unfixed plasmas (characters) of the racial type, for example, 
ought to be placed nearer to the fixed (and specific) plasmas (characters) 
and to be more adhesive to them than are the ancestral or individual plasmas 
(or characters). 

But on the other hand, living types are found (like Cyclops, Crepidula), 
where not only in the original egg and blastomeres, but even in some of the 
adult cells, paternal and maternal chromosomes remain distinct. Since, 
however, this singular, and yet unexplained fact of the longitudinal division 
of chromosomes at the beginning of each synapsis can be construed as a 
temporary separation of the maternal and paternal chromatin (or of these 
synthetic plasmas which represent and perpetuate them during the onto¬ 
genesis), we are led to conceive that a fertilized egg has the constitution 
shown in figure 1, where the reciprocal importance of each plasma is shown 
in relation to the formation of corresponding characters. 

But we must visualize these plasmas, with their probable constitution, 
which consists of elastic and malleable substance, flattened by reciprocal 
pressure. They ought then to be disposed towards each other, according to 
a definite order: so that the plasmas from each parent being of the same 
dynamic value and power, ought to accommodate themselves with those of 
the opposite side, just as the scales of a vegetable bulb are analyzed in 
layers. 


132 


LOUIS LEGRAND 


Thus figure 2 where the composite nucleus alone is shown, is more in 
accordance with facts than is the first one (fig. 1). 

With such a conception of hereditary substance by imbricated plasmatic 
“leaves,” of unequal value in transmitting various characters, we may 
consider what will be the result of their reciprocal coincidence and mobility. 



Schema. o[ Ike wkole Fertilise! Eg,§, 

Leaving for a moment these compulsory positions of each unfixed plasma, 
we believe that the possible reciprocal positions of these plasmatic planes 
are quite indefinite before amphimixis is accomplished. Just after the 
amphimixis shock is over, however, these plasmatic layers after hesitating 

































EXPLANATION OF HEREDITARY MECHANISM 


133 


for a moment are definitely fixed together, and remain so through all the 
subsequent karyokinesis during the life of the individual. 

The curves DC, DE, JE, JA (fig. 2), for example, are susceptible of in¬ 
definite variation, as the contact points D, F, H, G, are able to move 
both sidewise and forward or back. If this first design of plasmatic planes 
is exactly transmitted to blastomeres and to all the following cells of the 
body, as long as it is growing, there will be at our disposal a mechanical 
explanation of the slightest and finest hereditary transmissions throughout 
all the cellular generations in which ontogenesis is summed up, and we can 
understand that each unfixed character is able to maintain its individuality 
up to the end of individual growth. 

By this scheme, too, we avoid the problem of the transmission of fixed 
characters, because the corresponding plasmas are superabundantly sup¬ 
plied either by their continuity with the surrounding maternal ones, or by 
exterior autotropic digestive synthesis. 

If an explanation of latent characters is wanted, it is sufficient to admit 
that above all, the main point in order for a character to become manifested, 
in the figure as in reality, is not the mass of the plasmas present, but how 
much the surfaces of contact are stretched. This is then the degree of fixa¬ 
tion, of each of these plasmatic blocks, to their surrounding fixed plasmas. 

Consequently, only the zone which in the egg as well as in each cell, both 
unites and separates the two types of plasmas, is living and active. This 
zone of contact, of reciprocal sticking and fixation between cytoplasm and 
nucleus or nuclei can be called the zone of manifested characters. On the 
diagrammatic figure of the egg’s nucleus (fig. 2) this zone is indicated by the 
successive lines HA, AB, BC, CH; that is to say these plasmas, which the 
chance changes of amphimixis have brought to some central points like 
K, K', K", will have no hereditary action: They are not to be manifested 
as characters. Plasmas are not latent, but those characters which are 
dependent on them, are latent. 

Thus at the origin of each individual, some very important parts of 
their plasmas, which are effectively transmitted, have not, however, the 
least value in morphogenesis, because they are not affected by the zone of 
fixation. 

So can be solved the celebrated antinomy of a parent transmitting to his 
progeny some characters which he does not himself show while not trans¬ 
mitting many of his own characters. 


134 


LOUIS LEGRAND 


Ua[ixed PI asmas 


an 


A Characters. 



Paternal side 
Racial Plasmas 


.“Maternal Side 
Racial-Plasmas 

J&temal Side 
AneesUal.pl. 


Maternal Side 
Ancestral .PI. 

_Tat(ier’s 
Individual PI. 

Motfier’s 
Individual PI. 


Kg-2. 


Plasmatic Constitution oj* a Fertilized Egg’s Nucleus*. 

DOMINANT CHARACTERS 

These figures still afford an explanation why some types of unfixed 
characters are more likely to appear than are others: For example, the most 
frequent to appear are the racial ones, represented by the great surface 
HC + BC, and are evidently strengthened if both parents are born of the 






































































































EXPLANATION OF HEREDITARY MECHANISM 


135 


same race. It explains too, why some parental or maternal characters are 
dominant. This is for the same reason as that given above, namely, if their 
plasmas are flattening some portion of the manifestation zone, they are 
able to exclude, to eclipse those of the opposite sex: these eclipsed plasmas 
becoming unable to express themselves from the dynamic point of view, are 
called recessive. J 

The fertilized egg (fig. 2) shows the preponderance of the paternal racial 
characters, since CH is greater than CB; the exclusive appearance of some 
ancestral characters from the maternal lineage in AB, the corresponding 
paternal block being totally driven back inside the spheroid and therefore 
remaining eclipsed and recessive. Finally, such an egg nucleus shows also 
some imperceptible contact of parental characters (from paternal lineage), 
in H, and another, narrow zone of maternal origin, from H to A. 

The various phases of karyokinesis must be considered, by hypothesis, 
as tending to reproduce, in every new-born cell, the same plasmatic type. 
We can therefore conceive the repercussion of this type upon all organs and 
apparatus of the adult body and the automatic unrolling of resemblances 
proportionally with growth. All the body’s features depend on the way 
by which amphimixic chromatine combined. If the angle of meeting was 
slightly different, the whole form of the body would be consequently influ¬ 
enced and another hereditary type would be built. The geometrical com¬ 
binations of such lines and planes are quite innumerable, and we have to 
consider, not an explanation for all characters, but only for the unfixed 
ones, or more precisely, these morphologic modifications which the presence 
of these unfixed characters impress upon the total development of the fixed 
ones. 

In fact, the problem of heredity is greatly simplified, provided that our 
distinction between both plasmas and characters holds. Then, indeed the 
transmission of fixed characters and plasmas is an evident fact of material 
continuity, at each generation, by the nutritive reserves of the egg or the 
maternal blood. Their continuation during life is determined and assured 
by digestive assimilation, which rigorously and continually affords ade¬ 
quate supply of the fixed plasma’s material. 

These figures also may be used to explain the oldest, the most expressive 
of unfixed characters, sex. This governs all other unfixed characters and 
impresses something of its own nature. As it appears like some transition 
between both types, it ought to be inserted in the plasmatic relationships, 
between the big sphere of fixed plasmas and the nucleus of unfixed ones. 
It is then sufficient to admit, that, at the very moment of the amphimixic 
joining, when these curved planes are being accommodated to each other, 


136 


LOUIS LEGRAND 


sex is determined. This follows because the totality of one sex’s plasmas, 
when added together, must necessarily eclipse and exclude those of the op¬ 
posite sex. 

In other words, the sex which, in the total amount of the fixation surfaces 
of its representative plasmas in the specific sphere, has the greatest 



&>. 3 


of rnarufested Ckaracleta.nd 


Sexual J)omirtance 


amount of adherence,—this sex will literally dominate the other and make 
the corresponding characters prevail during the life of the individual: the 
other sex will then belong also, throughout life, to the series of latent 
characters before described. 

So the nucleus of figure 3 is destined to become female, since the curved 
fixation planes AB + AC are greater than CD -f- DB. However, this 






EXPLANATION OF HEREDITARY MECHANISM 


137 


female will wholly resemble her father since the adherent paternal plasmas 
in CD, have eclipsed or dominated the maternal ones, which have been 
repelled back inside the nuclear sphere. 

The first biologists who described karyokinesis (Strasburger, Fol, Biitschli 
Van Beneden, Hertwig) have attempted, in general, to attribute to it 
some relation to the transmission of character. Here we have attempted to 
define and limit this relation. It is enough to admit that the successive 
phases of karyokinese tend to maintain, in the new-born cells, the same type 
of accommodation and fixation of the unfixed plasmas both with each other 
and with the specific plasma (that is to say chromosomes), which was found 
in antecedent cells. 

These interactions between both sort of plasmas, their contacts, their 
mode of attraction and fixation, are therefore what constitutes life. 

The whole future individual has therefore its geometrical (design) re¬ 
corded and systematized inside that invisible protoplasmic sphere, which each 
histologist is able to fix, section and observe. All these figures ought to be 
considered as compulsory orientation planes, for future cell lineages. If an 
egg is to have its plasmas differentiated, as we have just considered in this 
study, it would not appear as a condensation, a reduced model of the whole 
organism, but rather as a sketch of the necessary scaffoldings along which 
the materials of that growing organism will come to form together and to be 
coordinated. 


SEX DETERMINATION IN ROTIFERS 


A. FRANKLIN SHULL 
University of Michigan , Ann Arbor, Michigan 

While there is some objection to applying the term sex determination to 
the alternative modes of reproduction in the rotifers, to do so has the advan¬ 
tage of recognizing a striking difference between the life cycle of the rotifers 
and those of most other animals which reproduce by the parthenogenetic 
and bisexual methods. In these latter more common forms, the sexual 
members of the cycle arise from parthenogenetic eggs which can develop in 
no other way, and from parents that are identical, or even from the same 
parents. The aphids and Cladocera are chiefly of this type. Even in the 
phylloxerans in which the sexual females and the males are derived from 
different parents, both develop from eggs that are necessarily parthenogene¬ 
tic. In the rotifers, on the contrary, while the males develop partheno- 
genetically, they are derived from eggs that may be fertilized. If fertilized, 
they yield females instead. The sex of rotifers is thus much more closely 
related to the mode of reproduction than in the aphids, Cladocera and phyl¬ 
loxerans. To describe as sex-determining those phenomena which lead 
to changes in the mode of reproduction in rotifers is a recognition of this 
more intimate relation. 

The factors leading to changes in the life cycle and to the determination 
of sex are various. Among the most important of these is heredity. This 
is demonstrated by the fact that different parthenogenetic lines employ the 
two modes of reproduction and produce the two sexes in very different 
ratios, even though reared under identical conditions. Crosses between 
lines exhibiting different ratios of male-production have been effected with 
results that can not for the present be reduced to any system. It is probable 
that a number of hereditary factors are involved in producing these 
differences. 

Though the hereditary factors concerned with the life cycle and deter¬ 
mination of sex are important—probably the most important—the extent 
to which they come to expression in any family may be greatly altered by 
environmental conditions. A long list of chemical substances has been 
found, each of which when dissolved in the water in which the rotifers live 

138 


SEX DETERMINATION IN ROTIFERS 


139 


reduces the proportion of sexual reproduction and hence of males. Some of 
these substances are ammonium carbamate, and other ammonium salts, 
cane sugar, butyric acid, and creatin. Concentrated solutions of some of 
these may practically exclude male-production even in lines which from 
hereditary factors alone produce many males. These substances are so 
unlike in their chemical properties as to throw no light on the cause of their 
common effect on the life cycle of the rotifers. Probably there is some 
unstable feature of the protoplasm which is readily disturbed by a variety 
of untoward conditions. 

Increase of male-production appears to be caused by very dilute solutions 
of certain substances (calcium chloride), but more particularly by the kind 
of food. It was first discovered by Whitney, and later verified in my own 
work, that green food, such unicellular organisms as Euglena and Chlamy- 
domonas, greatly increases the amount of male-production. Direct tests 
appear to show that part of this effect is due to the oxygen liberated in the 
water by the green organisms, but the major portion of it is evidently a 
phenomenon of nutrition, not of quantity but of kind. 

Another change in the amount of male-production in a given line of 
rotifers appears to result from long-continued parthenogenesis. Lines 
which produce many males in the early generations following the fertilized 
egg produce successively fewer of them in later parthenogenetic generations. 
Some biologists have attributed this well known change to an unsuspected 
change of environmental conditions, others to long subjection to uniform 
conditions, rather than to any internal phenomenon. Argument against 
the first suggestion is found in the fact that young lines introduced into the 
supposedly deteriorated conditions nevertheless produce many males. 
Against the second suggestion is the fact that hybrid lines from a cross 
known to result in high male-production produced many males under 
conditions in which similar hybrid lines produced long before were already 
showing a marked decline of male-production. In my opinion there is a 
cumulative change that is internal and largely independent of environment. 
Probably the progressive diminution of male-production is in no way due to 
parthenogenesis, but to the absence of bisexual reproduction. It might be 
more correctly described as due to long continued undisturbed metabolism, 
rather than long continued parthenogenesis. If this conception is correct, 
this progressive change would occur to the same extent in six months, 
regardless of whether only one generation or sixty generations had been 
produced in that time; and it is only because of the rapid succession of 
generations that the cumulative change can be expressed in a reduction in 
the number of males. 


140 


A. FRANKLIN SHULL 


None of these changes in the number of males produced appears to affect 
the hereditary nature of the line involved. When diminution of male- 
production is caused by substances in the water, removal of the unusual 
environmental condition immediately restores the male-production to its 
former high level. And when long absence of bisexual reproduction leads 
to a decline of male-production in a given line, the ability of that line to 
transmit capacity for male-production in crosses remains unimpaired. 

Crucial events in the determination of the mode of reproduction or of 
sex, in so far as they relate to a given individual, occur only in a very 
limited period of time. The mode of reproduction to be employed by any 
parthenogenetically produced female is fixed in a period not over two or 
three hours long, probably less, in the maturation of the egg from which she 
develops. The sex of any parthenogenetically produced individual is 
fixed in a corresponding short period in the maturation of the egg from which 
that individual’s mother develops. Since the critical part of the maturation 
of an egg occurs while the egg is still in the oviduct of the female, the sex of 
a male rotifer is determined in the body of its grandmother. Attempts to 
alter the mode of reproduction or the sex, by means of environmental 
conditions, before or after the maturation period just referred to, have 
invariably failed. 

The event in the maturation of an egg which determines whether the 
female developing from it shall produce rtiales or females is still unknown. 
Apparently there is no change in the chromosome number. Although 
males differ from females in the number of their chromosomes, there is no 
such difference between the mothers of males and mothers of females. 
There are twelve chromosomes in each of these types of female. Yet these 
females are irrevocably differentiated from one another by something. No 
female ever produces both males and females by parthenogenesis. 

What change occurs in maturation to differentiate the two types of egg 
so sharply from one another can only be conjectured. Probably it is a 
definite chemical change in the proteins of the egg. That change may have 
to do with the proteins of the chromosomes. A similar chemical change per¬ 
haps occurs in the germ cells of other animals in which sex is associated with 
chromosome number, as it appears to be in man; but presumably in such 
forms it is not the number that is important. If in such species the chemical 
change involves also a change in the number of chromosomes, while in the 
rotifers the chemical change leaves the chromosomes number unaltered, 
the two cases are brought under one point of view. Perhaps the nature 
of this chemical change, if such it is, will never be known; but in the search 
for it in the rotifers, it is not unlikely that the way is pointed by the fact 


SEX DETERMINATION IN ROTIFERS 


141 


that fertilized eggs all yield females. These eggs, without fertilization, 
produce males, and fertilization changes the sex as in the honey-bee. The 
essential chemical event, as far as sex is concerned, produced by fertilization 
may turn out to be the same as the event occurring in the maturation of the 
female-producing parthenogenetic egg. 

The above suggestions recall the metabolic theory of sex. The fact 
that the supposed chemical change in the rotifers occurs suddenly, that is, 
in a very short period of time, may make the chemical change theory 
seem incompatible with the general metabolic theory; for the metabolic 
theory usually assumes a gradual change, with a series of finely graded 
conditions between the two extremes which represent male and female. 
The incompatibility is not real, however. There may be in the rotifers a 
Series of metabolic processes of some other nature, or a variable metabolic 
condition, at some stage of which the sudden event in maturation occurs. 
I have avoided saying that the decisive event may occur at a certain 
“metabolic level,” because this expression has been used by those who have 
in mind some specific sort of change, such as the consumption of oxygen or 
the storage of fats or lecithin. It may well be that the chemical change 
that takes place in the rotifers can not be described in terms of “level.” 
However, although the current language of the metabolic theory may not be 
applicable to my conception of what occurs in the rotifers, it is apparent 
that there is nothing in the phenomena of sex determination in the rotifers, 
as far as these phenomena are known, which makes the metabolic theory 
of sex untenable. 


SOME DATA ON CONTROL OF SEX IN CLADOCERA 


ARTHUR M. BANTA and L. A. BROWN 
Carnegie Institution of Washington, Cold Spring Harbor, New York 

One of us has reared eight species of Cladocera under laboratory conditions 
for periods ranging from 11 to 118 months, from 39 to 448 generations (see 
table 1). In spite of the fact that reproduction has been solely by partheno¬ 
genesis, the stock has apparently lost nothing of its original vigor and there 
is an entire lack of indications of degeneration or of the occurrence of ab¬ 
normal individuals. Hence there is no evidence for an internal sexual cycle 
but on the contrary there is excellent ground for believing that partheno- 
genetic reproduction may be continued indefinitely and, by implication, 
that sex control is through external factors. 

Sex phenomena in Cladocera consist in the production of eggs requir¬ 
ing fertilization and of males. The parthenogenetic egg of Cladocera, 
which is the prevailing type, passes into the mother’s brood chamber and 
within about two days has developed into a freely swimming daphnid quite 
resembling the mother except for its small size. The sexual egg is much 
larger than the parthenogenetic egg. There are never more than two of the 
former in a clutch whereas there may be 12 to 40 of even more parthenoge¬ 
netic eggs in a single clutch. When sexual eggs are about to be produced the 
brood-chamber becomes covered by an ephippium, a heavy dense develop¬ 
ment of the carapace. This ephippium when cast off serves as a resistant 
case for the fertilized sexual eggs, which ordinarily develop only after a 
considerable dormant period. 

Circumstantial evidence that the sexual forms are called forth by environ¬ 
mental factors had been obtained from two sources before direct experi¬ 
mentation upon the subject was undertaken. It was noted that, whereas 
males and ephippial females did not often occur among our laboratory stock, 
when they did appear they were usually noted in more than a single strain. 
On one occasion many strains in the laboratory produced males within a 
comparatively short time. At another time males and sexual females were 
abundant among three species of Cladocera (Daphnia pulex and two species 
of Moina) in an outdoor pond from which our culture water was at that time 
obtained. Sexual forms appeared at the same time among several of our 
laboratory lines reared in water from this pond. 

142 


CONTROL OF SEX IN CLADOCERA 


143 


The first direct experimentation on the control of sex was with culture 
water from two different ponds—one pond containing males among its 
wild Cladocera population and the other lacking them. Males appeared 
with considerable frequency among the bottles made up with culture water 
from the pond which contained males among its wild Cladocera population; 
no males appeared in bottles containing culture water from the other pond. 
A second series of experiments consisted in rearing Cladocera in culture 

TABLE 1 


Periods of time and generations of descent of some laboratory lines of Cladocera 


SPECIES 

LINE 

NUMBER 

DATE CULTURE 
BEGUN 

NUMBER 

OF GENERA¬ 
TIONS TO 

9/21/21 

NUMBER OF 
MONTHS 


695+ 

11/17/11 

448 

118 


695- 

11/17/11 

417 

118 


713 — 

11/22/11 

429 

118 

Daphnia pulex. 

714+ 

11/22/11 

426 

118 


714— 

11/22/11 

437 

118 


984 

5/16/17 

193 

52 

D. longispina. 

898 

10/3/16 

220 

60 


795+ 

12/11/14 

297 

81 


795- 

12/11/14 

286 

81 

Simocephalus exspinosus. s 

740+ 

8/15/12 

378 

109 


740- 

8/15/12 

384 

109 

S. vetulus. 

1043 

10/28/20 

39 

11 

S. serrulatus. 

859 

9/28/15 

252 

72 

Moina affinis. 

851 

7/15/15 

382 

74 

M. macrocopa. 

1012 

5/27/18 

276 

40 

M. rectirostris. 

1010 

5/27/18 

248 

40 


water treated with precise amounts of NaOH and HC1. These experi¬ 
ments were frequently unsuccessful in bringing about the production of 
males, but when males occurred they appeared in bottles in which they had 
been anticipated,—and not elsewhere. 

The case then rested until the present collaborative work upon the prob¬ 
lem was undertaken. This work is still uncompleted and in progress but 
may be briefly reported upon here. 

It was found, as Grosvenor and Smith 1 had previously found for Moina 
rectirostris, that the simple expedient of crowding the females from early 

1 The life cycle of Moina rectirostris. Quart. Jour. Mic. Sci., 58, 1913, pp. 511-522. 























144 


ARTHUR M. BANTA AND L. A. BROWN 


life until they produce young causes the production of a proportion of males. 
Our procedure was as follows. Instead of only one (or two) females being 
reared in approximately 75 cc. of culture water, 10 or 20 females were reared 
in this quantity of culture water. Each experiment utilized the young of a 
single brood from one mother, hence any difference in sex of young in the 
crowded and uncrowded bottles is attributable to the effect of the crowding. 

The data for some of the lines tested are shown in table 2. The data 
for a given experiment are placed in horizontal lines. (Experiment 25 
was conducted in duplicate, hence there are two horizontal lines for it.) 
The totals and percentages of males by 1st, 2nd, and successive broods are 
placed below the tabulated data for a given line. Thus in experiments with 
Daphnia pulex Line 1005, Experiments 25 and 31, there were in the first 
broods of all “ten-mother” bottles 148 females and 22 males or 12.9 per 
cent males; in the second broods 171 females and 57 males or 25 per cent 
males; and in the third broods 64 females and 97 males or 60.2 per cent 
males; in all broods of ten-mother bottles for this line there were 385 females 
and 176 males or 31.3 per cent males. Data for Simocephalus exspinosus 
Line 795 plus, and for Moina macrocopa Line 1012 are also given in this 
table. 

The D. pulex data show that none of the six uncrowded mothers in three 
bottles produced any males in a total of 18 broods and 245 young; while the 
30 crowded mothers in three bottles produced 12.9 per cent, 25 per cent, 
and 60.2 per cent males, in their first, second and third broods respectively, 
or an average of 31.3 per cent males; and the 20 crowded mothers in a single 
bottle produced 0 per cent, 4.5 per cent, and 20 per cent males in the dif¬ 
ferent broods or an average of 6.4 per cent males. The S. exspinosus data 
show 0 per cenL males in two-mother bottles, 50 per cent males in a ten- 
mother bottle, and 68.2 per cent males in two twenty-mother bottles. The 
M. macrocopa data show 10.7 per cent males in the uncrowded bottles, 51.7 
per cent males in the ten-mother bottles, and 48.6 per cent males in the 
twenty-mother bottles. 

These are very limited data but they indicate that crowding is a ready 
means of bringing about the production of males. The fairly limited range 
of percentages of male young in the ten- and twenty-mother bottles of M. 
macrocopa suggests the possibility that in crowded bottles of this strain of 
M. macrocopa 50 per cent males is approximately the expectation. In the 
second brood of two two-mother bottles of Moina, and in the third brood of 
two other two-mother bottles of the same series males appeared. Either 
the factor calling forth males in crowded bottles may supervene to influence 
later broods in two-mother bottles or some other factor may also be opera- 


CONTROL OF SEX IN CLADOCERA 


145 


tive here. In the one experiment with S. exspinosus involving a ten-mother 
bottle no males appeared until in the second broods. The crowding effect 
seems not to have become manifest early enough to influence the first broods. 

The result common to the three species is 0 per cent or a low percentage 
of males in uncrowded bottles, 31 to 52 per cent males in ten-mother bottles 
and 6 to 68 per cent males in twenty-mother bottles. 

Similar experiments, some as successful, others less successful than those 
shown in table 2, have been tried with other strains of Cladocera. No strain 
at all adequately tested has failed to show some response to the effects of 
crowding (though in two strains the sexual manifestation has been in the 
production of ephippial eggs rather than males). Definite results have been 
obtained for Moina affinis, M. macrocopa, M. rectirostris, Daphnia pulex 
(three morphologically different forms of D. pulex have given positive re¬ 
sults), a species of Ceriodaphnia (one experiment only), Simocephalus 
vetulus, S. exspinosus and S. serrulatus. 

The specific environmental factor involved in the causation of males 
in crowded bottles is being sought. Formerly crowded bottles, with 
their recent population removed, when used again as culture bottles do not 
give males. Hence the cause of the crowding effect does not persist. The 
production of males is not due to mere scarcity of food since recently 
crowded or old depleted bottles give very few young yet the young are 
females just as those produced by uncrowded mothers with abundant food. 
The young produced by mothers reared in clear pond water, again with 
scarcity of food, are females. Hence age of culture medium, scarcity of 
food, or accumulation of excretory products (unless they are products of a 
very transitory nature) is apparently not the causative factor here. 

A single experiment suggested that the environmental factor involved 
may be the accumulation of carbon dioxide or depletion of oxygen in the 
crowded bottles. This will be tested further. 

Scanty data have been obtained bearing on the critical period during 
which the crowding effect must be operative. Table 3 presents some of the 
data bearing on this point. The data show that in their first broods none 
of the mothers in uncrowded bottles, or kept in crowded bottles less than 
about forty-eight hours, produced any males. From those kept crowded 
until fully forty-eight hours of age or older 45 per cent of the young were 
males, while those crowded all the time produced 34.1 per cent males. 2 The 

2 The higher percentage of males from those mothers crowded all the time than from 
those isolated between 48 and 76 hours is possibly not significant though it is possible 
there is a selective elimination of males under conditions of severe crowding. 

These data are not entirely satisfactory to the writers. The precise stages of develop¬ 
ment of the eggs were not known. In later experiments accurate observations will enable 
us to know whether the eggs have been laid or at approximately what stage of their ovarian 
development sex is irrevocably fixed. 


TABLE 2 

Data of some Cladocera sex control experiments 


146 


ARTHUR M. BANTA AND L. A. BROWN 


CROWDED BOTTLES 

Twenty mothers 

Fourth 

brood 

■b 



All twenty-mother bottles 




All twenty-mother bottles 


o 



Third 

brood 

r b 

IO 

t-H 

IO 

t-H 

b 

S3 

o 

CN1 

b 

£3 

NO 



b 

&3 

CNJ 

Co 

NO 

o 

09 

09 


Second 

brood 

b 



b 

IO 

b 

On 

t-H 

co 

O0 

b 

S3 

t-H 

X". 

LO 

b 

oo 

oo 

o 

85 

85 

NO 

NO 

First 

brood 

b 

o 

o 

b 

£3 

o 

276$ 

34 

46 

80 

b 

£3 

IO 

On 

NO 

o 

o 

CN 

t-H 

t-H 

CN 

t-H 

t-H 

OH t-H 

t-H CN 

35 

Ten mothers 

Fourth 

brood 

b 



All ten-mother bottles 




All ten-mother bottles 


o 



Third 

brood 

b 

(N tH 

NO CN •<-1 

r^. 

On 

b 

CN| 

o 

NO 

b 

CO 

—H 

<o 



b 

&3 

s 

o 

CN co On 
w (N (N 

64 


Second 

brood 

b 

N OO (N 
t-h t-h CN 

57 

b 

S3 

‘O 

CN) 

b 

NO 

t-H 

42 

42 

b 

£>3 

oo 

x-^ 

x^. 

b 

CN 

o 

O O N 
vO fO 

t-H 

T-H 

CN 

t-H 

CN 

T-H 

First 

brood 

b 

0 

22 

0 

22 

b 

S3 

On 

CN) 

385$ 

o 

O 

b 

S3 

o 

42$ 

o 

"O CO H* 

NO Tf CO 

OO 

T-H 

30 

30 

UNCROWDED BOTTLES 

Two mothers 

Fourth 

brood 

b 



All two-mother bottles 




All two-mother bottles 


o 



Third 

brood 

b 

o o o 

O 

b 

S3 

o 

b 

S3 

o 



b 

&3 

o 

o 

CO Cs ON 

tH tH CN 

t-H 

NO 


Second 

brood 

b 

o o o 

o 

b 

S3 

o 

b 

o 

o o 

o 

b 

&3 

o 

b 

o 

o 

NO 00 On 

CO co 

co 

CN) 

t-H 

63 

29 

92 

First 

brood 

b 

O O O 

O 

b 

&3 

o 

245$ 

o o 

o 

b 

£3 

o 

CH- 

On 

o 

CO O CO 
t-H CN CN 

T-H 

NO 

O r- 
co CN 

57 

aaawnN XNaKiaaaxa 

IO rH 

CN CO 

32 

33 


3 *0 
§■8 

*c*5 

« 


If) N 

P U 

C/3 l 

Q iO 
C On 


m 


CO - 



















































































































































































CONTROL OF SEX IN CLADOCERA 


43 

55 

98 

b 

© 

Ov 

LO 

All twenty-mother bottles 

! 

rH IO 

00 

© 

O O CN co 

t-h Ov 

t-H 

t". 

© 

CN 

b 

t-H 

b 

NO 

°o 

O © t-h CO 

M OO OO On 

O' 

O0 

CN 

© co t-h 00 00 

CO CO CO t-h 

o 

Ov 

rH 

b 

Ov 

O0 

H 

b 

Ov 

h ^ co © lo 

co io © 

OO 

Ov 

t-H 

H+ 1 © LO © © 

vo oo io o- 

rH rH 

Ov 

© 

b 

CN 

© 

LO 

o 

t-H 

CN 

© 

rH 

H H © co lo 

>0 N lO t-h t>» 

T-H T-H 

© 

© 

H 

VO IO 

Jt— CO 

rH 

rH 

b 

CO 

CN 

© 

All ten-mother bottles 


17 

50 

© 

© LO © t-H 

Tf IO 

CN 

© 

rH 

b 

© 

CO 

b 

ts. 

’N 

LO 

r)< N 

LO Ov CO CO 

© 

CN 

CN 

© -rf t-h >0 

CO T-H CO 
t-H 

© 

OO 

rH 

b 

© 

o^ 

LO 

b 

28 

© 

t-H © t-H O 

LO CO Hf 

CN 

CN 

t-H 

I s * rH O 

N 00 CO 

rH 

LO 

Ov 

CN 

b 

t— 

LO 

LO 

o 

co 

© 

O' © LO © 

O-J NO H 

rH 

H 

CO 

CN 

© © 

© 

b 

© 

All two-mother bottles 

b 

tN 

o 

"H 

© O' 

LO 

Cv 

LO 

© © © H H 
rH 

00 

rH 

b 

© 

rH 

O0 CO © OO © 

CO CO CN 

LO 

© 

rH 

© CN © © © 
CO 

Ov 

CO 

b 

H 

rH 

CN 

b 

LO 

ov h © © h © 

CN H N N CN 

CO 

©©©©©© 

© 

b 

© 

O 

© 

t— 

rf 

rj< O CN CO >0 >0 

CO LO Tf t-H t-H t-H 

Ov 

© 

rH 

21 

29 

30 

50 

51 
54 

Moina macrocopa 
(Line 1012) 















































































148 


ARTHUR M. BANTA AND L. A. BROWN 


TABLE 3 


Length of crowding and sex of young. Moina macrocopa (first broods only) 


EXPERIMENT NUMBER 


54 


65 


69 


70 


71 


74 


LENGTH OF CROWDING 


0 hrs. 

20-25 hrs. 

31-46 hrs. 

48-76 hrs. 

All time 
(72-96 hrs.) 

9 

cF 

% 
cF 

9 

cF 

% 

cF 

9 

^ ! cF 

9 

cF 

% 

cF 

9 

cF 

% 

cF 

15 

0 


15 

0 


17 

0 


0 

0 

21 

20 

19 

18 

0 

0 


155 

70 


15 

0 

0 

15 

0 

0 

17 

0 

0 

41 

37 

47.4 

155 

70 

31.1 

16 

19 

0 

0 







15 

16 
12 

0 

1 

0 


128 

0 


35 

0 

0 







43 

1 

2.2 

128 

0 

0 

15 

15 

0 

0 


12 

11 

0 

0 


8 

5 

7 

4 

0 

0 

0 

0 


0 

0 

4 

6 


44 

21 


30 

0 

0 

23 

0 

0 

24 

0 

0 

0 

10 

100.0 

44 

21 

32.3 

16 

15 

0 

0 


15 

13 

0 

0 





6 

0 

0 

11 


31 

39 


31 

0 

0 

28 

0 

0 




6 

11 

64.7 

31 

39 

55.7 

18 

17 

0 

0 


19 

20 

0 

0 





0 

15 

0 

5 

0 

0 

16 

0 

11 

3 

13 

8 


24 

101 


35 

0 

0 

39 

0 

0 




20 

51 

71.8 

24 

101 

80.8 

19 

16 

17 

19 

14 

0 

0 

0 

0 

0 


19 

20 

0 

0 


15 

16 
16 

0 

0 

0 





193 

31 


85 

° 

0 

39 

0 

0 

47 

0 

0 




193 

31 

13.8 




















































































































































CONTROL OF SEX IN CLADOCERA 


149 



critical period seems to be around forty-six or forty-eight hours,—about 
the time the eggs pass into the brood chamber. This critical period is 
not precisely enough localized as yet but these data suggest that the 
crowding condition must be operative until (or after) the eggs are laid and 
maturation is undergone. Hence it seems probable that in Cladqcera, sex 
is not irrevocably fixed until the maturation of the parthenogenetic egg, 
but that in a certain stage of the egg the sex of the forthcoming young 
is subject to control through environmental influences. 


















































DISTURBANCES IN MAMMALIAN DEVELOPMENT PRODUCED 
BY RADIUM EMANATION 

HALSEY J. BAGG 

From the Huntington Fund for Cancer Research , Memorial Hospital , and the Department of 
Anatomy , Cornell University Medical College , New York City 

The effect of radium on developing animals has been the subject of several 
researches since the early work of Bohn (1), in 1903, upon the ova and larvae 
of the sea-urchin. Experiments on developing nematodes, molluscs, 
amphibians, fishes, and birds are associated with the names of Perthes (2), 
P. Hertwig (3), Mottram (4), Tur (5), Schaper (6), O. Hertwig (7), G. 
Hertwig (8), and Bataillon (9). These investigators report the retarding 
effects produced by radiating the ova and developing embryos, showing the 
particular susceptibility of the nuclei of the cells, a general slowing up in the 
developmental processes, especially in the case of the central nervous 
system, and the total disturbances, depending upon the period of develop¬ 
ment when the radiation was applied, resulting in the formation of mon¬ 
strosities conforming more or less to a general type. 1 

Similar experiments concerning the effects of X-rays on development 
have been conducted by many investigators. After exposure to X-radia¬ 
tion, Perthes (10) and Hastings (11) noted abnormal cell division, and a 
retardation in the development of the ova of Ascaris megalocephala. Bor- 
dier (12), and Hastings, Beckton and Wedd (13) reported similar results 
following the irradiation of various insects. Gilman and Baetjer (14) after 
radiating the ova of Amblystoma, and Baldwin (15) the fertilized ova of 
frogs, were able to produce a fairly constant type of developmental defect. 

Injurious results have followed in all cases where mammals have been 
exposed to X-radiation. Forsterling (16), Langfellner (17), Krunkenberg 

1 In connection with the above statement, and applying to X-ray treatments as well, 
the question of dosage is an important one. A survey of the literature shows that there 
was a very wide range in the severity of the dose employed, and in several cases the 
experimental settings were inadequately described. (Bohn used “some centigrams” of 
pure radium bromide for from twenty minutes to two hours.) The amount of radium 
metal used in the investigations that have been mentioned varied from 2 mgs. to 35.1 mgs., 
and the time from a few.seconds to several hours. The deleterious changes in the animal 
tissues varied with the amount of radium and the time of exposure. 

150 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


151 


(18), and Walter (19) have shown that when any particular part of a young 
animal is exposed to a sufficient amount of radiation, that part fails to reach 
its normal size, and is unable to exercise a full degree of function. Lang- 
fellner (17) exposed pregnant guinea pigs to X-rays three days before term. 
The young, born a short time after treatment, were all dead. 

Arrests in development and the production of abnormal types may be 
induced not only by radio-activity, but by many physical or chemical agents. 
Abnormal temperature changes, treatment by many chemicals, lack of 
oxygen supply, or the overabundance of carbon dioxide, etc., have produced 
marked changes in the developing embryo. 

The present experiments are mainly concerned with disturbances in 
mammalian development, before and after birth, as a result of exposing the 
embryos of rats, at various times during the prenatal period, to irradiation 
from radium emanation. The effect on the embryos following radiation 
of the mother at varying intervals before mating was also determined. 
These experiments were designed not only to study the factors underlying 
the production of abnormal types, but through an examination of the 
abnormal to gain a clearer insight into the nature of normal development 
and differentiation. 2 


METHODS AND APPARATUS 

Two methods were used for applying the radium emanation. In the 
first method an “active deposit” was obtained by exposing a definite quan¬ 
tity of common salt to a comparatively large amount of radium emanation, 
about 500 millicuries were used, or the amount of radium emanation 
initially equivalent to 0.5 gram of radium metal. To the radio-active salt 
thus produced sufficient water was added to make a physiological solution. 
The pregnant rats were injected subcutaneously in the shoulder region, 
and intravenously through the caudal vein. Three to four minims con¬ 
stituted the usual dose. Because of the rapid loss of radio-activity of these 
solutions, the injections were made immediately after the preparation. The 
details involved in preparing and measuring the doses, as well as the methods 
for protecting the experimenter, are described elsewhere, (20) and (21). 
The activated solution exhibited all the known phenomena of radium metal 
itself, alpha, beta, and gamma rays were present, but the greatest physio- 

2 Dr. J. F. Gudernatsch was a co-investigator with the writer during the year 1919. 
A pre limin ary report of the work done with him at that time is given in the Proceedings 
of the Society for Experimental Biology and Medicine, 1920, xvii, 183. 


152 


HALSEY J. BAGG 


logical effects were probably due to alpha-ray activity. After long experi¬ 
mentation a dose of 5 millicuries was found to be the maximum applicable 
to the aims of this experiment. In the second method, gamma-ray radiation 
was applied through the ventral body wall of pregnant rats at nearly 
full term. A large amount of radium emanation was used, an amount 
equivalent to 1.5 grams of radium metal, filtered by 2 mm. of lead and 0.5 
mm. of silver. The source of emanation was 1 cm. away from the animal. 
The applicator, called a “lead tray” in clinical usage, was 6 cm. in diameter 
and 1.5 cm. high. This was placed in the bottom of a small wire cage, 10 
by 13 cm. in diameter and 10 cm. high, and was covered by a thin sheet of 
cardboard. The animal was placed on this paper immediately above the 
applicator. 

Preliminary tests showed that a dose of about 1300 millicurie hours was 
sufficient to produce developmental arrests in the embryos without killing 
the pregnant animals. Doses as high as 2900 millicurie hours, however, 
were successfully used in some cases. The embryos were killed by ether, 
and histological material procured at various periods after the treatment. 
The tissues were fixed in Bouin’s solution, cut in serial section, and stained 
with haematoxylin and eosin. 

EXPERIMENTAL RESULTS 

Series A. Injections of radio-active solutions 

1. Subcutaneous injections after mating. Sixty-five, full grown, normal, 
pregnant rats were treated in this series. They were divided into four 
groups, each treated at different periods after mating. Ten pregnant 
females were injected 7 days after mating; twenty-four, 10 to 14 days after, 
twenty-one, 15 to 17 days after, and ten, 18 to 21 days after mating. Many 
of the animals were killed at weekly intervals after treatment, although some 
were allowed to reach full term. 

Various degrees of developmental disturbances were noted, as shown in 
the following groups: 

1. There was a large number of cases where no embryos developed, or 
many began development and were early absorbed or aborted. The females 
in which no embryos were found, although they were definitely considered 
pregnant before treatment, occurred in the cases treated soon after mating, 
and in the instances where females were autopsied a considerable time after 
treatment. Figure 1 shows the remnants of maternal and embryonic struc¬ 
tures, and from the size of the placentae one can see that the fetuses had 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


153 


reached a fair degree of development before the radiation retarded the nor¬ 
mal physiological processes. In one case, see figure 2, a small ovoid sac 
was found attached to the uterine wall by a thin stalk. This appar¬ 
ently represented the remnants of a former embryo and placenta. Extrav- 
asated blood and cell detritus were found in this sac, and a great many 
large cells of an epithelioid nature that probably belonged to the former 
embryonic syncytium. The wall of this cyst was formed by fibrous con¬ 
nective tissue. 

2. Embryos were killed by the treatment, but were removed from the 
mother and preserved before they were absorbed. They showed various 
extravasations of the vessels of the subcutaneous connective tissue, within 
the meningial sinuses, and mainly along the dorsal mid-line of the body. 
Figure 3 shows a typical example of such a lesion which was situated in 
the mid-dorsal line. The mother of this embryo, no. 1167, was mated on 
April 22, 1919, injected with 4.9 millicuries on May 7, and was killed two 
days later. When the embryo was cut in serial section, it showed that the 
hematoma in the dorsal subcutaneous tissues had exerted sufficient pressure 
upon the spinal cord to produce at one place a complete dislocation. 
Microscopical examination of the viscera showed no pathological changes. 
Not all the fetuses of a litter were affected in the same degree. In one 
case 7 fetuses were found, 3 showing haemorrhagic lesions, 2 beginning 
to macerate, and 2 in the process of absorption. This difference in resist¬ 
ance was due either to the higher or lower vitality of the embryos themselves, 
or to the amount of radio-activity which passed the placentae. In another 
case the fetuses, although injured, were carried to full term, and among a 
litter of 6 young, 2 were apparentlynor mal and 4 showed haemorrhagic 
spots on the head, face, and along the dorsal mid-line of the body. 

3. Several young of a single litter showed areas of extravasation and 
were born alive. Their mother died, however, and foster mothers re¬ 
fused to nurse them. 

4. Eight litters gave normal living young. This number is low, because 
as previously stated, many pregnant rats were killed by the experimenter 
at various intervals after treatment. The average number of young 
per litter was 4.8, as compared with 6.5 per litter for the control rats, but 
the probable errors indicate that this difference is, most likely, not sig¬ 
nificant. Only one litter, containing 4 young, survived after a treatment 
given 7 days after mating. Several of the rats of this group, which had 
apparently escaped the full radium exposure during the uterine period, 
or perhaps they were more resistant to it, when mated inter se, produced 
litters of apparently normal young of normal fertility. The offspring of 


154 


HALSEY J. BAGG 


these animals, about 20 in number, were observed for two generations, but 
no abnormalities were noted. . 

2. Subcutaneous injections before mating. Seventy-seven females were 
treated in this group, 11 died as a result of the injection before they were 
mated, while several were killed at weekly intervals after mating, and 
some were allowed to continue to full term. Thirty-four animals were 
injected between 5 and 7 days before mating; seventeen, 10 to 14 days be¬ 
fore; and fifteen, 20 days before mating. 

Only three litters in this group showed abnormal young. The most in¬ 
teresting was a litter of 7, in which case the female was treated with 4.2 
millicuries, 22 days previous to fertilization, and the fetuses, approximately 
16 days old, showed very pronounced areas of extravasation, which in one 
case covered a large area on one side of the head and a few small scattered 
areas on the other side. These areas were not only along the dorsal mid¬ 
line, but also on the lateral surfaces of the body as well (see fig. 4). The 
lesions were much more widely distributed and more variable in size, than 
in the cases recorded under section 1. Although the conditions that 
produced these results were many times repeated, the above is the only case 
where positive data were obtained. Usually the female had either been 
rendered sterile, or the young were killed and absorbed at the early stages. 
There were two other cases, however, where young were found with 
haemorrhagic areas, and these occurred in a group of females that were 
treated 7 days before mating. Female 85 was given a dose of 6.6 millicuries 
on November 7,1919. It was mated on November 14, and as 3 young were 
bom December 11, fertilization took place about 14 days after the treat¬ 
ment. Two of the young were apparently normal, but one showed a large 
haemorrhagic area, which involved most of the right side of the snout, the 
right eye, and a portion of the lower jaw on that side. This area dis¬ 
appeared after three days. Female 99, injected and mated at the same time 
as female no. 85, received a dose of 5.6 millicuries. Five young, 3 males 
and 2 females, were born on December 13, making the date of fertilization 
about 16 days after treatment. One male and one female showed definite 
haemorrhagic areas on the face. Consideration of these cases will be 
deferred until later. 

Seventeen females that following treatment were killed at varying in¬ 
tervals after mating, showed markedly haemorrhagic or cystic ovaries, and 
the presence of much congested uteri. In these cases radium emanation 
apparently had either so altered the maternal tissues as to prevent 
fertilization, or development when started was soon followed by the death 
of the embryo and its absorption. Many nodules were found in the uteri 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


155 


in which it was impossible to differentiate between embryonic and maternal 
structures. 

The remaining females (as previously stated, 11 died between the period 
of treatment and mating) produced either full term normal young, or young 
apparently normal at autopsy. Several of these living young grew normally, 
were mated inter se, but produced no abnormal offspring although observed 
for two generations. 

3. Intravenous injections after mating. The intravenous injections were 
primarily planned to act as a check on the series of subcutaneous treatments. 
The object was to determine the immediate reactions that might occur in 
the embryo as a result of injecting a comparatively large dose of radio-active 
solution into the circulation of the pregnant female, and whether these 
reactions would be similar to those already recorded for the subcutaneous 
series. The toxic reactions were so prompt and fatal that it was not nec¬ 
essary to treat many animals to settle this point. A typical case is that of 
female no. 123. This animal, of about 19 days pregnancy, was treated with 
30 millicuries injected directly into the blood stream through the caudal 
vein. This was six times greater than the usual dose in the first two series. 
Three young were born dead 24 hours later. They showed very definite 
radium changes, typical of those already recorded for the subcutaneous 
series. A fetus was found still attached to an apparently normal placenta, 
but a characteristic area of extravasation was found over a considerable 
portion of the left side of the head. Another embryo showed two com¬ 
paratively small haemorrhagic areas along the dorsal mid-line, and the 
placenta in this case is also normal. The third fetus in this litter was 
apparently normal, but the placenta had acted in the nature of a shock 
absorber” in protecting the fetus from exposure to the radio-activity, and 
it was so swollen and completely filled with blood as a result of its injury, 
that it had the appearance of a large haemorrhagic sac. 

Series B. Results from radiating nearly full term pregnant rats 
with gamma-ray radiation 

Ten rats were treated at the end of about 19 days of pregnancy. It was 
found that exposure to about 1350 millicuries hours of radium emanation 
was sufficient to produce very decided changes in the embryo and yet leave 
the pregnant females sufficiently uninjured to be able to nurse their young 
and care for them until after the weaning period. When the dose was in¬ 
creased to 3378 millicurie hours the young were severely injured, and were 
either killed outright, or died 2 or 3 days after birth. 


156 


HALSEY J. BAGG 























DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


157 


The following are the conditions that resulted in the first generation of 
animals treated in utero with a dose of about 1350 millicurie hours. 

1. The young of each litter were born two or three days after the treat¬ 
ment, alive and apparently normal. 

2. About 10 days after treatment, about half of each litter became 
markedly anemic, showed symptoms of diffuse edema and promptly died. 
There was an easily recognizable slow development of meningial and 
spinal cord haemorrhages, similar to those already described as a result 
of treatment by radio-active solutions. In one case a young rat was found 
showing a typical haemorrhagic area in the region of the frontal lobes. The 
slow development of this lesion could be easily noted through the thin, 
transparent scalp. This young was one of several treated in utero with 
1350 millicurie hours of gamma-ray radiation on February 21, 1920. It 
was born 2 days later, and died on March 3. Another rat in this litter 


Fig. 1. Two Well-Developed Placentae Are Shown at the Left Attached to 
a Uterus Which Has Been Partly Opened 
The remnants of embryonic tissue are superimposed on the placentae. At the right 
is a placenta which has been dissected from the uterus, and shows more clearly the remains 
of embryonic material, here represented as a lighter area in the upper portion of the 
drawing. Female mated April 22, 1919, injected May 7, killed May 16. Dose = 4-6 
millicuries (subcutaneous). 

Fig. 2. A Stalked Sac Partly Dissected from the Uterus, Showing the Remnants 
of a Former Embryo and Placenta 

Female mated April 22, injected May 7, killed May 16. Dose = 4.8 millicuries 
(subcutaneous). 

Fig. 3. This is a Dorsal View of a Rat Embryo, Showing a Characteristic Area of 
Extravasation Due to the Treatment of the Mother During Pregnancy 
Female mated April 22, injected May 7, killed May 9, at which time seven fetuses 
were found about fifteen days in development. Two of the litter were macerated and 
two absorbed. Dose = 4.9 millicuries (subcutaneous). 

Fig. 4. Areas of Extravasation Are Shown in the Three Views of this Embryo, 
Similar to the Condition Shown in Figure 3 
But in this case resulting from treating the mother 22 days before fertilization. 
Note the wide distribution of the lesions. Female injected April 22, mated May 12,. 
killed May 30. Seven fetuses were found, 15 to 16 days old. Dose = 4.2 millicuries 
(subcutaneous). 

Fig. 5. There is An Extensive Meningial Extravasation Over a Considerable 
Portion of the Hemispheres, and a Hemorrhagic Lesion is Shown on the 
Reflected Skin from the Dorsal Interscapular Region 
This animal was treated in utero with 1350 me. hrs. of gamma-ray radiation on February 
21 and was born apparently normal on February 23. It died on March 2. 



158 


HALSEY J. BAGG 


shows the presence of three distinct hemorrhagic areas, a small frontal 
lesion, a fairly extensive one in the occipital region, and a small lesion in the 
subcutaneous tissues in the thoracic region, near the mid-dorsal line on the 
left side of the body. This animal also died on March 3. A third animal 
belonging to the same litter is shown in figure 5. Here is seen a still more 
acute reaction, as shown by the fact that the animal died a day sooner than 
in the two cases above, and there is an extensive area of meningeal hemor¬ 
rhage which covers most of the dorsal portion of the brain, involving the 
frontal and occipital regions and the medial area between, as well as a 
considerable portion of the right temporal area. In addition, a distinct, 
rounded hemorrhagic lesion may be noted on the reflected skin on the left 
side of the body. This lesion occurred in the mid-shoulder region of the 
back. 

The heads of several of the young rats showed marked lateral compression. 
In one case a hemorrhage so affected the spinal column as to produce com¬ 
plete paraplegia. The tissues of these animals were studied histologically. 
Save for the mechanical disturbances produced by the presence of the 
extravasated areas, the most marked pathological conditions were seen in 
the liver and intestines. In the first case there was a pronounced fatty de¬ 
generation of the hepatic cells, and in the second, a desquamation of the 
lining cells of the intestinal mucosa. 

3. It is interesting to note that the other half of each litter survived the 
treatment, grew to a normal size, and some animals have lived for over 18 
months. They showed the effects of the late uterine treatment by the 
following arrests in development: 

a. The first pathological condition that was noted was that the eyes 
became smaller, the pupils opaque, and there finally was a complete, or 
nearly complete closing of the lids and total blindness. This condition 
was first observed a short time after the eyes had opened. In one animal, 
which had grown to normal size and weight for its age, the left eye was 
nearly completely closed, both pupils were opaque, but the right eyelids 
were slightly more opened than those of the other side. The animal was 
one of a litter treated in utero on March 8,1920, was born six days later, and 
these observations were made on March 1, 1921. The dose in this case 
was 2920 millicurie hours of gamma-ray radiation, which was a dose higher 
than one usually tolerated. 

b. Mating tests showed that both the males and females were completely 
sterile in the first lots, but subsequently a first generation female, that 
had been treated with 1350 millcurie hours, mated with a male similarly 
treated and gave birth to 9 apparently normal young. 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 159 

c. Before these adult offspring of treated animals were killed for histo¬ 
logical examination, their neurological reactions were very carefully 
studied. The animals, being blind, when startled assumed various 
defensive attitudes, but save for these reactions their behavior was remark¬ 
ably normal. There was no ataxia in locomotion or in any of the feeding 
reactions, auditory acuity was normal, and there was no cutaneous hy- 
poesthesia, or other sensory disturbances. Except for blindness, there was 
nothing to suggest abnormal sensory function. 

d. When these animals were autopsied, marked developmental dis¬ 
turbances were noted in the condition of the central nervous system. The 
cerebral hemispheres were greatly reduced in size, and in several cases very 
little cortical material remained. Those portions of the brain that were 
ontogenetically older (the archiostriatum and the cerebellum) were appar¬ 
ently normal. The optic tracts were markedly atrophic. Correlated with 
this disturbance in brain development, the skull was found to be asymmetri¬ 
cal, narrow, thicker than normal, and concave in the frontal region. 

One animal, which belonged to the same litter as those of section 2 of this 
series, was treated with 1350 millicurie hours on February 21, 1920, was 
born on February 23, and was killed December 31, 1920. This was one of 
the animals which (except for blindness) showed no abnormal neurological 
reactions. A dorsal view of the brain shows an apparently normal cere¬ 
bellum and normal olfactory lobes, but the part of the brain which repre¬ 
sents the rudiments of the hemispheres shows a great lack of development 
of cortical substances. A side view of the brain shows that the cortex is 
very thin indeed, not completely covered what should normally be the 
frontal, occipital and lateral aspects of the brain. The remains of the 
hemispheres do not sufficiently approach each other in the median line to 
cover the colliculi beneath. It was possible in this specimen to see the 
lateral ventricles through the transparent membranes. Several other 
brains have been studied which showed various degrees of developmental 
arrests resulting from radium treatment. In some cases the hemispheres 
were markedly reduced in size, were widely divergent in the median line 
and yet the pallium was complete over the entire surface. In all these 
cases there was marked optic atrophy. These brains are now being sec¬ 
tioned, and a study of them in greater detail will be the subject of a separate 
communication. 

e. A histological study of the eye showed that the eyeball was reduced to 
one fourth the normal diameter. The retina was missing but traces of the 
choroid remained as a few scattered pigmental cells. The cornea was three 
times as thick as the normal and covered with four or five layers of opaque 


160 


HALSEY J. BAGG 


squamons epithelium. The optic nerve was extremely small, not more 
than one third the normal dimensions. 

/. The testes of the radiated animals were decidedly atrophic, and a 
comparison with the normal is shown in the photograph in figure 6. The 
diameters of the testicle alone (minus the epididymis) of the experimental 



Fig. 6. At the right is shown a normal untreated testicle of a white rat surmounted by a 
well developed epididymis, which is partly obscured by fat. At the left is a radiated testis 
showing considerable atrophy. The single small lobe at the very bottom of the photo¬ 
graph represents the remains of the tail of the epididymis, the head and body of that 
part being completely missing. The animal was treated in utero on February 21, 1920, 
was born February 23, and was killed December 31, 1920. Dose = 1350 millicurie hours 
of gamma-ray irradiation. 

animal was 14 mm. for the length and 7 mm. for the width, while the control 
measurements from normal animals of the same age and weight and with 
the same method of fixation were 21 mm. for the length and 11.5 mm. for 
the width. The epididymis of the radiated testis was practically missing. 
A small portion of the tail remained, but the head and body of the epididy- 



DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


161 


mis had failed to develop. Histological examination shows that there is 
little evidence of spermatogenesis. Some tubules seem to contain imperfect 
spermatoblasts and forming spermatozoa, but the great majority of tubules 
show complete degeneration and loss of epithelial cells, and contain loose 
granular material, which in places is calcified. Some spermatic tubules are 
greatly dilated and filled with granular material. Very few interstitial 
cells are visible. 

The ovary of the radiated animals was reduced to one-fourth or one-fifth 
the normal size. The Graffian follicles were entirely missing. Groups of 
lutein cells persisted in small numbers, but showed marked hydropic de¬ 
generation. Some of the large vessels about the ovary were sclerosed. 

g. The liver, kidney, lungs, spleen, and the other organs were examined, 
but showed no pathological disturbance. 

CONTROL GROUP 

Pregnant rats of the same stock, the same age and weight, were injected 
subcutaneously and intravenously with equal amounts of solutions that 
previously had been strongly radio-active, but were allowed to “decay,” 
until they had lost their radio-activity. These experiments gave absolutely 
negative results. As a control to the gamma-ray experiments pregnant 
rats, sisters of the treated animals, were allowed to breed under exactly the 
same experimental conditions. No abnormal young were observed. 

DISCUSSION AND SUMMARY OF RESULTS 

It has been shown that when doses of radio-active solutions are injected 
into an animal marked physiological reactions take place. Large doses 
produce severe toxemia, resulting in pronounced pathological changes in 
the various viscera of the white rat (20). A study of metabolic changes in 
dogs, as determined by urine analysis, showed that following intravenous 
injections of such solution, there were very decided increases in the total 
nitrogen content of the urine, the urea, creatinine, uric acid and the total 
phosphates (22). A prompt reduction occurred in the number of white 
blood-cells of the dog after intravenous injection of these solutions, 
associated with a marked decrease in the relative percentage of circulating 
lymphocytes (23). In order to reduce as much as possible the severity of 
reaction, very small doses of radio-activity were used in the experiments 
recorded in this article. But even with comparatively small doses, certain 
rats treated in utero showed very acute reactions. Many were killed 
by the treatment and were absorbed or aborted. Others were found 


162 


HALSEY J. BAGG 


showing pronounced areas of subcutaneous extravasations, mainly situated 
along the mid-dorsal line of the body and within the meningial sinuses. 
This condition was probably due to the destructive action of radium on the 
endothelium of the blood vessels, as well as a possible increase in blood pres¬ 
sure, as was shown to occur in the dog by Burton-Opitz and Meyer (24) 
after intravenous injections of very small quantities of radium bromide. 
A similar reaction of the blood vessels to radiation was previously reported 
by Halkin (25) for the skin of pigs, by Danysz (26) for radiated mice, and 
later, was again confirmed by Obersteiner (27). This destructive action of 
radium on the blood vessels is in line with clinical observations on the usual 
prompt regression of very vascular tumors (the angiomata, in particular) 
after exposure to irradiation. 

The changes in the rat embryos of this experiment are interesting in so 
far as they show that a sufficient amount of radio-activity was able to pass 
the placenta and subsequently affect the developing embryo. This occurred 
after subcutaneous as well as intravenous injections of the mother. By far 
the most interesting observation concerned the presence of lesions similar to 
those described above in rat embryos whose mother was treated with radio¬ 
active solutions a considerable time before mating. The writer has no 
explanation to account for this phenomenon. It would appear that the 
treatment of the mother several days previous to conception has lessened 
the faculty of the later-developing embryo to form proper endothelium of the 
blood vessels, and the wide distribution of these lesions over the body of the 
embryo (peculiar to this group of animals) would tend to substantiate this 
view. One female was injected 22 days before fertilization, and since the 
solutions lose their radio-activity very rapidly (there is about a 50 per cent 
reduction in the first hour after the preparation) the likelihood of any 
radio-activity remaining over during this period and affecting the egg at a 
later critical moment is remote. The amount of radio-activity remaining 
after 22 days, if present at all, should, as determined from physical com¬ 
putation, be infinitesimally small. 

The series of intravenous injections again emphasize the specific action 
of radium emanation in the production of typical areas of subcutaneous 
extravasations in the developing young, and in addition shows that the 
placenta may act in the nature of a “shock absorber” and prevent the 
embryo from receiving the full effect of the radiation. 

We now come to a consideration of the cases wherein pregnant females 
were treated with external applications of comparatively large doses of 
gamma-ray radiation. At this time a report is given only for embryos 
treated towards the end of pregnancy. The writer plans to continue this 
line of investigation and treat at earlier prenatal periods. 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


163 


The results emphasize the well known delayed reaction associated with 
gamma-ray radiation. There was approximately a ten-day interval follow¬ 
ing treatment during which no changes were noted in the embryo, and 
during this period the young animals were born in an apparently normal 
condition. Acute reactions promptly occurred at the completion of this 
time, killing half of each litter. The young rats died showing typical radium 
changes, such as, anemia, diffuse edema and meningial, spinal cord, and 
subcutaneous extravasations. These extravasations were markedly similar 
to those already described for the series of solution treatments. The liver 
in these animals showed a fatty degeneration of the hepatic cells similar to 
the condition reported by Mills (28) after exposing a series of mice to gamma- 
ray radiation. The only other pathological change noted in these embryos 
was a desquamation of the lining cells of the intestinal mucosa. This 
observation is in line with the results emphasized by Hall and Whipple (29) 
in their experiments on roentgen-ray intoxication in dogs. 

While the animals described above died after showing acute reactions, 
certain of their litter mates (half of the litter) continued to develop appar¬ 
ently normally. This difference in reaction may possibly be due to indi¬ 
vidual variability or tolerance for the radiation, but it most likely can be 
explained by the fact that certain embryos were probably slightly farther 
away from the source of radiation than others, and as the intensity of 
radiation varies inversely as the square of the distance, even such slight 
differences in distances that did exist would be sufficient to subject the 
embryos to a considerable range in intensity of radiation. This is especially 
important in this case because the source of radiation was only one centi¬ 
meter from the body wall of the mother. The quality of radiation, how¬ 
ever, remained the same for all the embryos. 

It was soon apparent that the animals that lived over the ten day period 
had not completely escaped the effect of the radiation, as was shown by a 
suppression of the full development of the eyes. Eye defects were noted, 
such as opaqueness of the pupil, atrophy of the lens and closing of the lids, 
which resulted in complete blindness. These animals grew to a normal 
size, successfully competed with their cage mates for food, and showed 
absolutely no abnormal neurological condition, except those clearly incident 
to blindness. At autopsy, in some cases over a year after birth, very decided 
developmental arrests were noted in the structure of the brain. All grades 
of such maldevelopment were noted in the condition of the neopallium, 
from merely a decrease in the size of the hemispheres, which permitted the 
corpora quadrigemina to be clearly visible from above, to a more marked 
atrophy of the cerebral cortex until only a very thin lamina of tissue re- 


164 


HALSEY J. BAGG 


mained to represent that structure, and there were large areas in the 
frontal, occipital, and temporal region where no cortex existed at all, so that 
when the meninges were removed the basal ganglia were clearly seen from 
without. 

This correlation between defects in the development of the eye and the 
brain has been emphasized by Stockard (30) in his recent paper on develop¬ 
mental rate and structural expression. He states as follows: “The periods 
of arrest necessary to induce the eye and the brain modifications are so 
close together or so nearly the same, that one generally finds combina¬ 
tions and mixtures of the defects among the same experimental group of 
embryos.” Again in the same article Stockard has shown that the type 
of deformity that results from experimental disturbance depends upon the 
developmental moment at which the interruption occurs. It is significant 
that the animals of this experiment showed arrests in the development of 
the neopallial portions of the brain and not in those regions which are onto- 
genically older. Apparently the radium emanation, acting towards the end 
of pregnancy, had affected the development of the brain after the basal 
ganglia, the cerebellum and medulla had become fairly well differentiated, 
and therefore these portions showed no gross changes. But the radium had 
slowed the developmental rate of the neopallium (which we know is one of 
the last portions of the brain to differentiate) during its period of active cell 
proliferation, and that portion of the brain was never able to reestablish 
its proper rate of development in relation to the other parts of the brain. 
If the period of treatment had occurred earlier in prenatal existence, other 
portions of the brain would probably have shown disturbances as well. 
The writer does not believe that the deformities in the brains of these 
animals were due to the early production of vascular disturbances later 
recovered from, but to an actual inhibitory effect of the radiation upon the 
developing nerve cells. If extravasations had occurred in this group of 
animals, and were so situated as to affect the development of the cerebral 
cortex in particular, they probably would have been detected as were even 
the comparatively small lesions which were associated with the acute 
reactions. Also, if the effect was largely due to vascular disturbances, 
from the nature of the radiation employed, one would expect more general¬ 
ized changes throughout the entire brain. However, on the other hand, 
Craigie (31), in his recent paper on the relative vascularity of various 
parts of the central nervous system of the albino rat, suggests that the 
vascularization of the more recently evolved centers (of the cortex cerebri) 
is more susceptible than the more ancient regions to sexual, hereditary or 
environmental influences. ” 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


165 


From a neurological point of view, it is interesting to consider that the 
animals with practically no cerebral cortex reacted so normally in their 
ordinary behavior. Except for blindness there was no other apparent 
sensory disturbance, and motor coordination appeared perfect. The physi¬ 
ological functions localized in the cerebral cortex of the rat were in these 
animals apparently transferred to the basal ganglia and other paleokinetic 
portions of the brain, showing the remarkable degree of compensation 
possible to the mammalian brain when the disturbing element acts at an 
early period in its development. In this connection it is worth mentioning 
that the radium emanation did not produce a sudden traumatic effect, as is 
normally the case with the experimental production of brain lesions, and, in 
fact, the radium changes were probably prolonged over a considerable 
period. This condition favored the establishment of compensatory reac¬ 
tions, and exists, (as shown by the writer in a recent article (32) ) even in 
the case of radium lesions experimentally produced in adult mammalian 
brains. 

The reproductive system completes its development a considerable time 
after birth, and so it is not at all surprising that these structures should have 
shown a considerable amount of atrophy due to the developmental arrest 
during the prenatal period. The ovaries and testes appeared to suffer with 
equal severity. 

An interesting correlative relation was shown by the fact that the other 
viscera (digestive, excretory, etc.) of the animals that showed marked 
developmental arrests of the nervous and reproductive systems were ap¬ 
parently normal, and the animals grew to an average size. The lungs, 
liver, kidneys, etc. had differentiated before the physical agent was 
employed. Further studies with earlier prenatal treatments should throw 
some light on establishing the critical growth periods of the various em¬ 
bryonic structures. 

As a final point of interest it is worth considering that the results of this 
investigation may be of interest to the clinicians and the laboratory workers 
who handle large quantities of radium and utilize X-rays. Although the 
results of this paper deal only with irradiation of the female, there is no 
reason to believe that the germ cells of the male are more resistant to these 
destructive agents than those of the female, and, in fact, there is very 
good experimental evidences to show that spermatozoa of some animals 
are expecially likely to produce abnormal young after exposure to compara¬ 
tively small quantities of irradiation. Physicians should guard against 
the possibility of producing developmental arrests such as shown in this 


166 


HALSEY J. BAGG 


article when treating pregnant women, as well as the possibility of altering 
the human germ cells by irradiation previous to conception. 3 

Again, the writer wishes to call the attention of those interested in the 
sterilization by irradiation of the unfit for eugenical purposes, to the need 
of extensive experimental work on the larger mammals before any such 
measure be extensively used. 


CONCLUSIONS 

1. The marked selective action of radium emanation on fast growing 
embryonic structures was noted in these experiments. 

2. Very decided developmental arrests occurred in the differentiation 
of the nervous and reproductive systems of mammalian embryos exposed 
to irradiation towards the end of pregnancy. 

3. Experimental animals with greatly reduced, or practically no neo¬ 
pallium, gave apparently normal neurological behavior except for blindness. 

4. Radium emanation, used either in the form of a radio-active solution 
injected into the adult female, or employed as an external gamma-ray 
radiation, produced marked areas of extravasation in the subcutaneous 
connective tissue of the developing young. This suggests that the action 
of radium emanation might be selective upon the endothelium of blood 
vessels. 

5. Extravasations occurred in the developing young of females treated 
with radio-active solutions a considerable time before fertilization, and 
suggest that in some way the faculty of the later developing embryos to 
form proper blood vascular endothelium had been interfered with. 

6. The results so far obtained indicate that gamma-ray radiation is a 
physical agent admirably adapted to the study of experimentally produced 
developmental arrests in mammalian embryos. 

7. When women are subjected to therapeutic irradiation, especially 
during the early stages of pregnancy, the clinician should be forewarned 
concerning the possibility of producing very grave disturbances in the 
developing child. 

ACKNOWLEDGMENTS 

The writer acknowledges with pleasure his indebtedness to Dr. James 
Ewing for his aid in the interpretation of the pathological results. 

8 The writer does not mean to be understood as stating that present day clinical irradia¬ 
tion treatments produce such effects in the developing young, but it is his personal opinion 
that such changes are biologically possible. It is not possible to obtain desired information 
by comparing the amount of exposure that a small mammal can stand with the corre¬ 
sponding dose that a man should tolerate judging by comparative weights. The small 
mammal can tolerate very much more radiation in proportion to its weight than a man can. 


DISTURBANCES IN DEVELOPMENT PRODUCED BY RADIUM 


167 


REFERENCES 

(1) Bohn: Compt. Rend. Soc. de biol., 1903, xl, 1442. 

(2) Perthes : Deut. med. woch., 1904, xxx, 632. 

(3) Hertwig, P.: Archiv. f. mikros. Anat., 1911, lxxvii, 301; ibid., 1915-16, lxxxvii, 63. 

(4) Mottram: Archiv. Middlesex Hosp., 1913, 12th Can. Rep. 

(5) Tur: Arch, de Zoo. exp6r. et g6n6r., 1906, v, Ser. 4, 39. 

(6) Schaper: Deut. Med. woch., 1904,1434. 

(7) Hertwig, O.: Archiv. f. mikros. Anat., 1911, lxxvii, no. 2,1. 

(8) Hertwig, G.: Archiv. f. mikros. Anat., 1911, lxxvii, 165. 

(9) Bataillon: Rev. g£n. des Sciences pures et appliq., 1911, xxii, 786. 

(10) Perthes: Deut. med. woch., 1904, xxx. 

(11) Hastings: Archiv. Middlesex Hosp., 1912,11th Can. Rep. 

(12) Bordier: Le Radium, 1905, xi, 410. 

(13) Hastings, Beckton and Wedd: Archiv. Middlesex Hosp., 1912, 11th Can. Rep. 

(14) Gilman and Baetjer: Amer. Jour. Physiol., 1904, x, 222. 

(15) Baldwin: Anat. Rec., 1919, xvii, no. 3,135. 

(16) Forsterling: Verhand. d. deut. Ront. Gesell., Ill, 126; ibid, v, 68. 

(17) Langfellner: Miinch med. woch., 1906, 44. 

(18) Krunkenberg: Verhand. d. deut. Ront. Gesell., v, 70. 

(19) Walter: Fortschritte a. d. geb. der Ront., 1912, xix, 123. 

(20) Bagg: Jour. Cancer Res., 1920, v, 1. 

(21) Duane: Boston Med. and Surg. Jour., 1917, clxxvii, 787. 

(22) Theis and Bagg: Jour. Biol. Chem., 1920, xli, 525. 

(23) Bagg: Jour. Cancer Res., 1920, v, 301. 

(24) Burton-Optiz and Meyer: Jour. Exp. Med., 1906, viii, 245. 

(25) Halkin: Arch. f. Dermat. u. Syphilis, 1903, lxv, 201. 

(26) Danysz: Compt. rend. Soc. de biol., 1903, cxxxvi, 461; ibid., 1903, cxxxvii, 1296. 

(27) Obersteiner: Arbeiten a. d. Neurolog. Inst. Wien, 1905, xii, 96. 

(28) Mills: Lancet, 1910, ii, 462. 

(29) Hall and Whipple: Amer. Jour. Med. Sci., 1919, clvii, 453. 

(30) Stockard: Amer. Jour. Anat., 1921, xxviii, 115. 

(31) Craigie: Jour. Comp. Neur., 1920, xxxi, 429. 

(32) Bagg: Amer. Jour. Roent., 1921, viii, 536. 


THE GENETIC SIGNIFICANCE OF THE ALCOHOLIC TREAT¬ 
MENT OF WHITE RATS 

E. CARLETON MacDOWELL 
Station for Experimental Evolution , Long Island, New York 

Even today the lay mind pursues us with the question, “Have you 
decided yet whether nature or nurture is the great factor? ” Yet it is clear 
that no simple answer is possible for this question. The real problem is not 
a general comparison of the two factors, but a study of the way each factor 
operates. After making this study, specific cases may be analyzed. If it 
is a common error to expect a final victory for either nature or nurture, it is a 
much more common error to expect a single, simple result from experiments 
involving alcohol. How far the specific findings herewith reported for white 
rats may apply to different animals is a matter for experiment, not con¬ 
jecture; but if one generalization may be permitted, it must be that the 
action of alcohol on animals is complex, it works through more than one 
channel, and the end results are due to a balancing of various tendencies. 
Instead of being on the plane of adding hydrochloric acid to sodium hydroxid 
and obtaining sodium chlorid and water, experiments with alcohol are more 
nearly parallel in the complexity of the results, to dropping a bomb upon 
a whole chemical laboratory. Recognizing the complexity of a problem is 
one step towards its solution. 

The following discussion is based upon experiments carried on for five 
years at the Cold Spring Harbor Station for Experimental Evolution. 
Data are at hand showing that alcohol, administered by the inhalation 
method, tends to modify the treated white rats in three ways: (1) by lower¬ 
ing their growth rate, (2) by reducing their fecundity, (3) by retarding 
their success in habit formation. The main question under investigation 
may be stated as follows: does this alcohol treatment have any genetic 
significance; is the race as well as the individual modified in any way by this 
treatment? The present experiments appear to give an affirmative 
answer—the alcohol has modified the race. But just how this has been 
brought about and how to appraise the final results can only be suggested 
at present. Opposing influences are working: the untreated children and 
grandchildren of the treated rats produce litters smaller by 10 per cent 
than their respective controls, and they tend to be less successful in learning 

168 


ALCOHOLIC TREATMENT OE WHITE RATS 


169 


the true path through the maze; yet they grow faster than their controls 
and they produce from 30 to 50 per cent more litters in the same time. 
Clearly the effects of the original alcohol treatment are shown in the follow : 
ing generations, but with equal clarity does it appear that these effects are 
of different sorts. 

Primarily it is a matter of finding through what channels alcohol works 
and not of judging how bad are the results. For to know the modus 
operandi of alcohol is fundamental, while the final results may differ in 
animals or man according to different superimposed influences. In the 
rats the criterion of habit formation appears to give evidence that the 
alcohol has directly modified the germinal material so that a difference in 
behavior can be detected; this appears in the untreated children and grand¬ 
children, in the treated offspring from treated parents, and in the untreated 
offspring from treated parents and treated grandparents. A change has 
occurred that is not removed by the absence of the alcohol, nor is it increased 
by the application of alcohol to a second generation. But there is no 
evidence that this modification is located in the chromosomes, or that this 
will continue to appear through an indefinite number of generations. Litter 
size gives evidence of the same type of germinal modification—a 10 per cent 
reduction throughout all generations. Upon growth and the number 
of litters alcohol must act in a different way. It is probable that the 
alcohol has not produced a modification at all, as far as these characters are 
concerned, but rather has eliminated germinal material that bore weaker 
producing power and slower growth. For, in spite of the marked reductions 
found in the treated animals, their untreated descendants were superior to 
their controls in both these regards. Thus there is evidence that alcohol 
may work in two different ways in the same animals, it may directly change 
germinal material and it may act as a selective agent. Stockard is about to 
present evidence of the modifying effect in guinea pigs; Pearl has found 
evidence of the selective action in fowl. It is then not a case of attempting 
to reconcile these two results or to decide which is correct; the results are 
different and both are correct. The two results emphasize two different 
channels of alcohol action, which, as shown, may occur simultaneously 
in the same animals. In both of these ways may the race be influenced: 
it is genetically significant that alcohol may act as a modifier of germinal 
material; it is genetically significant that alcohol may act as a selective 
agent between existing germinal differences. 


INHERITANCE IN MENTAL DISORDERS 
HENRY A. COTTON 

Medical Director of the New Jersey State Hospital at Trenton 

The entire subject of the role of inheritance in mental disorders has been 
developed largely from a non-biological standpoint, and it was not until 
the important stimulus of the work of the Eugenics Record Office, under 
the able direction of Dr. Charles B. Davenport, began to be reflected in the 
work of various state institutions that any scientific data was obtainable. 
For years, even centuries, “insanity in the family” was considered the most 
important factor in the production of a psychosis; later, many of the so- 
called “hereditary” types were found to be due, largely, if not entirely, to 
extrinsic factors, and as the importance of these extrinsic factors was dem¬ 
onstrated the hereditary factors lost their importance. 

For instance, in paresis, when it was found that the invasion of the brain 
by the organism of syphilis was the principal, if not the only, cause of the 
disease, it was not logical or accurate to assume that “mental heredity” 
could be the cause of the invasion of the brain by the germ of syphilis. 
Hence, paresis in adults has been quite properly eliminated as a “hereditary 
disease” although the juvenile type is due to an inheritance of the syphilis 
germ, or, rather, to a direct transmission of this organism from the mother 
to the child. 

The same theory applies to organic brain diseases in general, such as 
brain tumor, arteriosclerosis, etc. Those interested in the subject of hered¬ 
ity are quite willing to admit that these disorders depend very little upon the 
inherited factors. 

With the so-called functional mental disorders, the lack of any known 
extrinsic factors in their causation led one to believe that the inherited 
factors were most important. This viewpoint was further fortified by the 
fact that in at least 70 per cent of such disorders heredity was present. As 
none, except psychogenic factors, were found, it was apparently obvious 
that the inherited constitution and the unstable personality were responsi¬ 
ble for the breaking down of the individual under mental stress. In 
other words, we were still dealing in these conditions with disorders of the 
mind and not 6f the brain. 


170 


INHERITANCE IN MENTAL DISORDERS 


171 


It is interesting to note that Davenport and others are trying to account 
for the inheritance of these conditions by inherited disturbances of the 
endocrin system, a step certainly in the right direction, although as yet no 
definite proof has been given. 

From the study of modern biology we learn what is inherited and what 
is not. Conklin is very positive in his statements which follow: 

Developed characters, whether acquired or not, are never transmitted by heredity and 
hereditary constitution of the germ is not changed by changes in such characters. Possi¬ 
bly environmental stimuli, acting upon germ cells at an early stage in their development, 
may rarely cause changes in their hereditary constitution, but changes produced in 
somatic cells do not cause corresponding changes in the hereditary constitution of the 
germ cells. Germ cells, like somatic cells, may undergo modifications which are not 
hereditary; if starved they may produce stunted individuals, and this effect may last for 
two or three generations. They may be stained by fat stains and the generation to which 
the) r give rise be similarly stained; they may be poisoned by alcohol or modified by 
temperature and such influence may be carried over to the next generation without 
becoming hereditary. Environment may profoundly modify individual development, 
but it does not generally modify heredity. 

With these biological tenets before us we certainly should be very cautious 
in ascribing to heredity the most important r6le in the production of the 
“functional” psychoses. 

As has been stated, the theory of the role of heredity in mental disorders 
was developed largely through statistics dependent upon state hospital 
records, and anyone who has examined such records is impressed with their 
inaccuracy and errors. Heredity meant “insanity in the family” whether 
in the direct or collateral lines, and no investigation was made as to the 
nature of the psychoses, whether somatic, paretic, etc. From this belief 
we have happily been led by Davenport to more accurate studies in 
this field. 

At the State Hospital in Trenton an intensive study was made of the 
hereditary factors in mental disorders, and the investigation was continued 
for a period of five years. Some 50,000 individuals were charged and as 
was the case in other hospitals many families were found afflicted with in¬ 
sanity. On the other hand a number of cases, large enough to be conspicu¬ 
ous, were found in which there was no hereditary taint. These so-called 
negative families seemed to me to be more important that the positive 
ones. 

Further, after carefully studying the material, no laws could be deduced, 
no inheritance of definite type, but rather a mixed and confusing array of 
facts which offered little help in solving the problem of the causation of the 
functional mental disorders. In spite of the obvious evidence in some cases 


172 


HENRY A. COTTON 


of the inherited character of these disorders, in a large number of instances 
the negative cases would loom up in a disconcerting manner and weaken our 
confidence in the importance of heredity. 

In the past five years intensive studies made at the State Hospital in Tren¬ 
ton have convinced me that in these so-called “functional” disorders cer¬ 
tain extrinsic factors, formerly overlooked, are of far more importance than 
the hereditary factors in producing the psychosis. These studies have also 
confirmed the modem biological axiom that function is directly related to 
stmcture. 

If this be true, and there is no one today who would doubt such a fact, 
then abnormal function must be accompanied by abnormal structure. 
Applying this to the psychoses we are justified in concluding that we are 
dealing with disorders of the brain, rather than disorders of the mind, in¬ 
dependent of the brain. The converse of this law still has adherents, but 
scientific fact will sooner or later replace such opinions. They are the 
relic of the belief that the mind was independent of the brain, and such 
beliefs are contrary to modern biological teaching. 

If we have destroyed the role of heredity in the production of the 
psychoses, what are the factors responsible for mental disorders? By 
utilizing the methods developed in modern medicine we have been able to 
demonstrate that the so-called “functional” psychoses were accompanied 
by multiple foci of chronic infections of which the patient was often entirely 
ignorant, and because of the hidden or masked character of these infections 
they were entirely overlooked by the physician. This was particularly 
true of the psychotic patient, for heredity and psychogenic factors were 
considered the sole agents responsible for the mental disorder. 

The doctrine of the role of heredity was necessarily fatalistic for if the 
patient were doomed at birth to have a psychosis why attempt to prevent 
such a calamity and why make an effort to look for other causes. That 
such an attitude has existed cannot be denied and it has certainly stifled 
investigation in purely medical fields. However, it has not hindered the 
prosecution of studies in the psycho-analytic field. 

It has been found that one of the principal causes of these mental disorders 
was cerebral toxemia arising from the toxemia produced by chronic foci 
of infection found in the teeth, tonsils, gastro-intestinal tract, genito-urinary 
tract, etc. I am not prepared to say just what the mechanism is in these 
cases, whether the poison acts directly upon the brain or indirectly through 
the endocrin system, but I am of the opinion that with or without heredity 
these mental disturbances will supervene if the toxin is sufficiently virulent 
and the individual lacks resistance to the toxin. 


INHERITANCE IN MENTAL DISORDERS 


173 


It is not my purpose to enter into a discussion of this work except to 
say that we find that patients suffering from the so-called “functional” 
mental disorders often have multiple foci of infection and, further, that 
when these infections are eradicated in the early stages of the disease prac¬ 
tically all will recover. As a proof of this statement I will give the results 
of our work at the State Hospital for the last three years. The spontane¬ 
ous recoveries for a period of ten years prior to 1918, averaged 37 percent. 
In the last three years this recovery rate has been doubled. Further, out 
of 380 patients in this group, admitted in 1918, only 50 remain in the hos¬ 
pital today and 9 of these are criminals. A recent survey of these cases by 
fieldworkers has shown that the cases considered recovered are, after three 
years, perfectly normal and the number of recoveries is larger by ten than 
at the time of discharge. By that we mean patients discharged im¬ 
proved, who recovered after leaving the hospital. 

What place are we then to give heredity in the constellation of causative 
factors? It certainly cannot be absolutely ignored, but we can still evaluate 
its importance without giving it the exalted position it formerly occupied. 

Does the toxin circulating in the blood of these patients alter or damage 
the germ plasm, rendering the individual more susceptible to this germ late 
in life? Are congenital abnormalities of the colon, occasionally found in 
parent and child, inherited characteristics which predispose to infection late 
in life, or is the infection carried by the parent transmitted to the child 
in infancy by direct contact, as in tuberculosis, and does it through years of 
development produce a psychosis late in life? 

These questions are yet to be answered. I believe, however, that the 
theory of direct inheritance in mental disorders has little to support it at 
present. As it does not affect the prognosis of a given case its discussion at 
this time is merely academic. 

These views do not in any way diminish the importance of eugenics, but 
rather open up a broader scope for eugenics in studying how the human race 
can be benefited. Such studies should not be limited to heredity alone. 


COMPARISON OF THE INCIDENCE OF THE SUPRACONDYLOID 
PROCESS IN GROUPS WITH NORMAL AND 
ABNORMAL MENTALITY 


R. J. TERRY and LEE D. CADY 
Washington University , St. Louis , Missouri 

Discrepancy in the incidence of the supracondyloid process as found in 
different nationalities and in the insane and criminal classes has given rise 
to the conjecture that, assuming the methods of computation to be compara¬ 
ble, the cause may lie in the physical differences in the different races and 
classes which have furnished the figures. Thus Wenzel Gruber (1) in St. 
Petersburg stated the incidence as 2.7 per cent, whereas Testut (2) in 
Lyons found the frequency to be only 1 per cent; the studies in each case 
having been made in the dissecting laboratory. Those who have examined 
the humeri of colored races have expressed the belief that the variation is 
much rarer than among white peoples, but there no figures for accurate 
comparison. Nicolas (3) in France, found the process in 6 individuals out 
of 115 insane subjects, and calculating the incidence on the total number of 
processes (bilaterality was present in three cases) reached the high percent¬ 
age of 7.8. A still greater frequency is claimed for Italian criminals by 
Ferdinando (4) who has given the incidence as 11.29 per cent. 

Recently two independent investigations on the incidence of the supra¬ 
condyloid process have been carried on by the writers, one concerned with a 
group of people assumed to be normal mentally, the other dealing with the 
patients of an insane asylum. The results indicate a slightly greater fre¬ 
quency of the variation in the second group but the acceptance of such a 
conclusion generally is not yet justified, as will appear from the following 
account of the subjects and methods involved in the calculations. 

The study of the so-called normal group was made in the year 1920 (5). 
One thousand patients in the Dispensary of the Washington University 
School of Medicine taken at random were examined for the presence of the 
supracondyloid process by the method of palpation. The group consisted 
of 683 white subjects of which 316 were males and 368 females; 515 were 
adults and 168 under age; the remaining 317 were negroes of which 120 
were males, 197 females; 248 were 21 years and over and 69 were under age. 

174 


INCIDENCE OF THE SUPRACONDYLOID PROCESS 


175 


In regard to the diagnosis, although there were psychiatric patients in the 
group none was defined as insane; the great majority came to the clinic on 
account of some physical ailment incidental to all people. The group is a 
fair example of the physical and mental status of the poorer class of a large 
city, suitable for a study in the living of a variation of which our knowl¬ 
edge has been derived from investigation of cadavers of the dissecting 
laboratory. 

The second group was chosen on the basis of mental defect of the subject 
and consisted of one thousand individuals. These were all patients of the 
St. Louis Sanitarium, a public institution for the care of the insane. The 
group was composed entirely of adult white patients, 500 males and 500 
females. Regarding the medical diagnoses, an extensive range of mental 
disorders was included in the examination for the variation. 

The method employed for the discovery of the supracondyloid process in 
the living is very simple: the examiner seated, palpates with the fingers of 
one hand the arm of the subject who stands before him, while with the other 
hand grasping the patient’s forearm he induces slight passive motion. 
Recognition of the presence of a process will be favored by its prominence 
and by the leanness and supplenness of the patient’s arm. Processes less than 
4 mm. in height are found with difficulty even under favorable conditions 
and undoubtedly larger processes would be missed in fat or muscular arms. 
The earlier investigations of the supracondyloid process have afforded 
abundant evidence that the high development of muscles has no relation to 
the origin and size of the bony spur, for it may be present at birth, occurs 
commonly in women as well as in men, and in the left arm apparently as 
often as in the right. Moreover some of the largest processes have been 
observed in subjects of frail build and conversely tiny spurs present them¬ 
selves in robust, muscular arms. When a process was discovered by the 
method of palpation, X-ray pictures were made to verify the observation 
and to permit of more detailed study than is possible by the manual ex¬ 
ploration, by this means the form, size and location of the apophysis can be 
determined very accurately. 

The value of the combined method of palpation and X-ray had been 
estimated by comparing the results both in incidence and physical charac¬ 
ters, with those gotten by observation of the skeleton and cadaver in the 
anatomical laboratory. Thus the occurrence of the variation in the mixed 
group of the Dispensary was represented by 7 persons in 1000; Testut 
found the variation in 8 subjects out of 929 in his laboratory. 

Whereas in the entire dispensary group but 7 individuals presented the 
supracondyloid variation, in the 1000 insane patients examined at the 


176 


R. J. TERRY AND LEE D. CADY 


Sanitarium, 12 bore the process as revealed by palpation and the X-ray; 
an incidence of 1.2 per cent as against 0.7 per cent for the normal group. 
This difference for the entire ground will be somewhat lessened when the 
comparison is made between only white subjects; the figures would then be 
1.2 vs. 0.878 per cent. If the comparison be limited to adult whites, then 
the difference will very nearly disappear (1.2 vs. 1.16 per cent); but such 
restriction is not called for in view of the fact already mentioned, that the 
variation is well known in children even to the new-born. 

The mode of distribution relative to sex and symmetry cannot be known 
until far greater numbers of authenticated data are at hand. There is con¬ 
siderable evidence of the process occurring with about the same frequency in 
both sexes and in both arms. In the dispensary group there were 4 males 
and 3 females having variation, whereas in the group of insane there was 1 
male and 11 females exhibiting the process. No general conclusion should 
be drawn from these figures or from tabulations of the distribution with 
respect to symmetry which in the group at large was 6 for the left and 2 for 
the right; in the insane 6 left and 7 right. 

Diagnosis of the insane patients having the variation revealed one instance 
each of dementia, hysterical insanity, paranoia and senile dementia; two 
cases of simple melancholia; two of manic depressive insanity; four instances 
of dementia praecox. From this it appears that no correlation of the proc¬ 
ess with a particular type of insanity obtains. 

Investigation of inheritance of the variation is in progress for both groups. 
At the present moment three instances of the process have been found in 
one family of the dispensary group . 

The group from the dispensary has been used for comparison with that of 
the insane asylum not because it was considered best suited to the purpose 
but because it comprises the only sort of subjects on which observations 
have been made and for which figures are available. The intelligent and 
physically fit element of the population is the last to become available for 
studies of this nature. However the question raised by the high incidence 
of the process in the insane given by Nicolas cannot be considered to be 
answered (even though his group is too small for a satisfactory calculation) 
until the incidence has been found in a selected number of people vigorous 
in mind and body. The results of the present comparison certainly do not 
lend support to the idea of marked frequency of the process in the insane: 
they indicate only slightly a higher incidence in the insane we have examined. 


INCIDENCE OF THE SUPRACONDYLOID PROCESS 


177 


REFERENCES 

(1) Gruber, W.: Monographic des Canalis supracondyloideus humeri. Mem. Acad. Sci. 

St. Petersburg, 1859, iii. 

(2) Testut, L.: L’Apophyse sus-epitrochleene chez l’homme. Intemat. Monatschr. f. 

Anat. u. Physiol., 1889, vi. 

(3) Nicolas, A.: Nouvelles observations d’apophyse sus-epitrochleene chez l’homme. 

Rev. Biol, du Nord de la France, 1890-91, iii. 

(4) Ferdinando, U.: Processo sopracondiloideo dell’ omero. Parma, 1899. 

(5) Terry, R. J. : A study of the supracondyloid process in the living. Am. Jour. Phys. 

Anthrop., 1921, iv. 


THE RELATION OF HEREDITY TO TUBERCULOSIS 

s 

PAUL A. LEWIS 

From the Henry Phipps Institute of the TJniversity of Pennsylvania , Philadelphia 

Notwithstanding this subject is very old, one may hope to add some new 
evidence bearing upon it and to indicate that such conclusions as the pro¬ 
fession may possibly have come to are properly subject to revision. 

From the beginning of medical time it was considered that tuberculosis 
ran in families; that heredity had much to do with the occurrence of the 
disease. With the discovery of the tubercle bacillus and the realization 
that tuberculosis was due to an infectious agent, it came to be believed that 
probably hereditary influences were more apparent than real, and that in¬ 
dividuals in certain families more often had tuberculosis, not because of 
any special influence of blood or breeding, but because the disease having 
been established in that family, the chances for contagion and contact were 
much increased, and all facts could be explained in the light of our knowledge 
of tuberculosis as an infectious disease. 

Since the establishment of the infectious nature of tuberculosis, a certain 
amount of work has indicated that possibly there was something more to the 
matter and that the older view could not be lightly dismissed. 

A number of years ago Dr. Pope went over all the material then available 
at the Saranac Lake Sanatorium, making a tabulation of this material on 
the basis of relationships of patients, in order to find out whether husband or 
wife were more liable to contract tuberculosis from each other than were the 
children of tuberculous parents. This material was collected and taken to 
England and put into the hands of Pearson who was interested in the study 
of this kind of material. Pearson applied mathematical methods to the 
study and seemed to show to his own satisfaction that there was evidence 
of hereditary influence on the development of tuberculosis in this material, 
namely, there was certain evidence that the marital relatives contracted 
tuberculosis in a larger degree than the average population considered 
without reference to the incidence of tuberculosis in the marriages. 

There was also a larger influence evidently due to the blood relationship; 
that is, the children of tuberculous parents were not only more liable than 
the average population to have tuberculosis, but were more liable to have 
tuberculosis than the husbands of tuberculous wives or the wives of tubercu- 

178 


RELATION OF HEREDITY TO TUBERCULOSIS 


179 


lous husbands. There seemed to him a satisfactory evidence that blood 
had something to do with it. He reasoned that the chances of infection were 
as great between husband and wife as they were between parent and child. 
These results of Pearson’s, however, have not been generally accepted by 
medical men up to date, and there are two reasons why this is so. First, 
from the fact that very few medical men were able to understand the method 
by which the results were arrived at and the terminology in which they were 
expressed. The second reason was biological, based on the proposition that 
most medical men thought that not sufficient account had been taken of the 
various susceptibilities to tuberculosis as they were here involved, and the 
fact was disregarded that children were more apt to be infected than adults, 
and this greater susceptibility to infection during childhood might ade¬ 
quately serve to explain the figures. 

A pupil of Pearson’s (Goring) undertook to restudy the same sort of 
material by taking prisoners in a certain English prison as the basis, and 
again it was found that children of tuberculous prisoners had tuberculosis 
in an appreciably larger percentage than the children of non-tuberculous 
prisoners. There was no difference discovered between the wives or hus¬ 
bands, as the case might be, of tuberculous prisoners as against the non- 
tuberculous. It was reasoned, then, that this had answered the objection 
cited in a fairly convincing way, because the prisoners are not much in con¬ 
tact with their families, either the marital relatives or the blood relatives, 
and there was an appreciable difference in favor of the proposition that the 
incidence of tuberculosis was higher when blood lines were followed. It is 
difficult to say what impression was made by the work of Pearson and Gor¬ 
ing and what the prevailing opinion is in such a matter. Suffice it to say, 
the influence of heredity as predisposing to tuberculosis has been quite 
largely discounted by the members of the medical profession and by others 
whose opinion carried weight in public health work. 

A great deal of work done in our laboratory has led us to the conviction 
that within any species of animal there is a large variation in resistance to 
tuberculosis. Rats and mice, which as species are very resistant, show a 
very wide individual variation also. Guinea pigs and rabbits as species are 
much less resistant, but also show a wide range of individual variation. 
The records of experiments with cattle indicate similar individual variation, 
and from what we know of tuberculosis in the human species, we may sur¬ 
mise that there are racial differences, but we may assume with greater as¬ 
surance that individual susceptibility varies greatly. 

We have tried to determine by experiment whether these individual 
variations had any true basis in heredity. The work has been made possible 


180 


PAUL A. LEWIS 


by a cooperative arrangement with the Bureau of Animal Industry of the 
United States Department of Agriculture. Dr. Sewall Wright, of the 
Bureau’s staff, is jointly associated with me in all that follows. 

Some fifteen years ago the Bureau of Animal Industry at Washington 
studied certain problems in the field of genetics and heredity on the basis of 
experiments on guinea pigs. The problem they studied was: What is the 
influence of inbreeding on the general qualities of the animal? Does 
inbreeding, as many stock raisers maintain, inevitably cause deterioration, 
or does it not? To satisfy themselves on this point, they had started from 
twenty-three pairs of guinea pigs from which they had continuously bred 
by a very rigid system of brother and sister mating. The answer which 
they found was that inbreeding did not, of and by itself, cause deterioration. 
The primary effect is a fixation of character, good, bad, or indifferent. A 
different combination became fixed in each family. Families with an es¬ 
pecially unfortunate combination did not long survive. 

We have used five strains which had been retained from those older ex¬ 
periments to test resistance to tuberculosis. The system of brother-sister 
mating has been long enough in operation so that the strains are practically 
homozygous. They are found to differ by as much as 100 per cent in their 
length of life after standard inoculation with B. Tuberculosis. 

The results suggest that the hereditary influence is exerted through at 
least three and possibly four factors or groups of factors capable of being 
transmitted separately. 

It would certainly throw much light on the fundamental nature of natural 
immunity to tuberculosis if the physiological characteristics of these hy¬ 
pothetical factors could be determined. Our efforts in this direction have 
not as yet led to any decisive result. Dr. Wright’s study of the Bureau 
records has shown that such attributes as age, weight, ability to gain weight, 
fertility, and ability to bring to maturity young born alive are, even 
when considered together, of slight importance as related to resistance to 
tuberculosis. 

It may possibly turn out that more significance is to be ascribed to the 
suggestion which a recent experiment has developed, namely, that these 
guinea pig strains appear to differ in their reaction to a local tuberculous 
infection of the skin and in their reaction to certain inflammatory irritants, 
such as croton oil and cantharides. We have not so far been able to formu¬ 
late an estimate of these differences in terms of resistance factors, and the 
matter is referred to only to show the trend of our further search for 
explanatory evidence. 


RELATION OF HEREDITY TO TUBERCULOSIS 


181 


How far is it useful or proper to go in the application of the experimental 
results so far obtained, to an interpretation of human tuberculosis? It is 
now generally regarded as true by biologists that wherever factors or groups 
of factors can be transmitted as units in heredity in one species of either 
plant or animal, those factors will be similarly under hereditaiy influences 
wherever they may occur. It is also recognized that when such factors are 
found in one species, they are very likely to be common to other related 
species as among mammals for example. There is, however, no a priori 
assurance that the relative importance of the factors will be maintained 
from species to species. There is, then, reason to expect that the factors 
influencing resistance in the guinea pig are also a human possession; until 
we can definitely recognize these factors as such and follow them into the 
human species by observation, we will be unable to dogmatize on their 
effectiveness as influences governing our racial reaction to tuberculous 
infection. 


THE INHERITANCE OF CANCER IN MICE 


LEO LOEB 

From the Department of Comparative Pathology of Washington University, St. Louis 

We studied the significance of heredity in the etiology of cancer through 
the long continued observation of a considerable number of strains and 
families of mice which were kept under identical external conditions, but 
were bred separately. Our investigations were begun in a preliminary 
way in 1907 and continued, with the aid of Miss Lathrop, on a much 
larger scale from 1910 on. Altogether we kept the records of approximately 
12,000 female mice which were observed throughout the whole period of 
their life. The strains were followed through consecutive generations. 

The following is a brief summary of our principal conclusions: 

1. The cancerrate of each strain or family is a definite characteristic of 
this strain and is transmitted by heredity to successive generations. The 
differences in the Tumorrate in various strains are very pronounced; the 
Tumorrate may vary between zero in certain strains and almost 100 per 
cent in others. 

2. These differences in rate persist with a surprising regularity through 
successive generations in the majority of strains. In certain strains, how¬ 
ever, variations in the Tumorrate do occur. Usually these changes consist 
in a decrease in the Tumorrate in later generations; they are in all proba¬ 
bility due to two factors: 

a. In certain families and strains as a result of long continued inbreeding 
a gradual decrease in fertility and vigor occurs. Associated with this 
change is, in certain cases, a noticeable decrease in the Tumorrate. 

b. Various factors cause a selection to take place within the strains; 
certain families die out, while others gain preponderance. This change is 
accompanied in certain cases by a change in Tumorrate. 

3. If we cross strains with a similar Tumorrate, the offspring inherits the 
Tumorrate common to both parents; if both parents differ in Tumorrates, 
the Tumorrate of the offspring is on the whole intermediate between those 
of the parents. But all degrees of intermediacy are observed. In our 
material the number of strains in which the rate of the parent with the higher 
Tumor-incidence dominated was on the whole greater than the contrary 
one. 


182 


INHERITANCE OF CANCER IN MICE 


183 


4. The age at which tumors appear is just as characteristic of individual 
strains as the Tumorrate. The Tumorage is also transmitted by heredity. 
We can distinguish two factors in the inheritance of the Tumorage: 

a. In general in the strains with the higher Tumorrate the Tumors appear 
at an earlier period of life than in the lower Tumorrate strains. Both 
Tumorrate and Tumorage are due to the action of an intensity factor. The 
greater this intensity, the more numerous are the individuals affected and 
the earlier the Tumors appear in these individuals. 

b. In addition to this general intensity factor which applies to many 
strains, there is a peculiar Tumorage in certain strains which is independent 
of the Tumorrate. Strains with a similar Tumorrate may have a different 
Tumorage and in hybrids Tumorrate and age may be inherited separately 
in the offspring. 

5. In general the cancerrate in mice is not a sex-linked character. This 
fact does however not exclude the possibility that in certain cases a sex- 
linked factor may enter as one of the multiple factors which determine the 
inheritance of cancer. Certain of our observations suggest such a possi¬ 
bility. 

6. Our investigations make it possible to express in a quantitatively 
definite manner the hereditary tendency to cancer in individual strains of 
mice, the figures varying in different strains between zero and 100. This 
hereditary tendency is, however, not a simple quantity but composite, 
because, ( a ) The hereditary disposition to cancer is due to the cooperation 
of multiple factors. The results of hybridization suggest this interpretation. 

(b) There is hidden in this figure a second factor which again is variable: 
namely, the activity of the ovary. In all the strains the realization of the 
hereditary tendency to cancer presupposes the activity of the internal se¬ 
cretion of the ovary. Without this cooperation no cancer can originate. 
With the full activity of this factor the hereditarily transmitted intensity 
character determines the upper limit of the cancerrate. Again the intensity 
of this ovarian factor can be expressed in a quantitative manner, the quan¬ 
tity in this case representing the time during which the ovarian internal 
secretion had a chance to act. If this secretion is eliminated in mice in the 
early stages of adult, sexually mature life, mammary cancer is practically 
prevented from appearing even in normally high tumorrate strains. The 
longer the ovarian function had a chance to act, the more the cancerrate 
increases up to the range which is given in the figure for the hereditary 
tendency to cancer. Suspension of breeding also usually diminishes some¬ 
what the cancerrate, but to a very much less extent than the exclusion of 
the internal secretion of the ovary which latter is the true realizing factor, 


184 


LEO LOEB 


the cooperation of which is necessary. Injury to the mamilla by suckling 
young can therefore not be an important factor in the causation of mammary 
cancer in mice. 

7. Thus it has become possible to express in a quantitative way the tend¬ 
ency to a disease (cancer) as due to the interaction of two main factors, 
both internal, the one hereditarily fixed and the other accessible to experi¬ 
mental variation. Both factors combined are the predisposition as well as 
the cause of cancer. In the case of other kinds of cancer, conditions are 
presumably similar, the role of the ovarian function being probably taken 
over in such cases by other glands with internal secretion or by external 
stimulation. 

8. Is it possible to associate the hereditary tendency to cancer with other 
factors active in the life of mice? No definite causal connection between 
tendency to cancer and such characters as vigor, prolificity, size, color can 
be found. Connections which may be found are probably not causals 
but due to coincidence. These other characteristics are inherited just as 
the tendency to cancer, but are in crosses distributed among the hybrids 
independently of the predisposition to cancer. 

9. There may however possibly be an exception to this independence of 
the hereditary transmission of the tendency to cancer. If we determine 
among various strains of mice the mortality or age of death from all other 
causes except cancer, we find that the high cancerrate strains are very 
largely strains in which the tendency to die from other causes than cancer 
at an early period of life is especially pronounced. This connection is pre¬ 
sumably not due to coincident but to a causal relationship. 

10. The tendency to die from other causes than cancer at a certain period 
of life is also hereditarily transmitted, but it varies among different strains 
much less than the predisposition to cancer. Again the tendency to die 
from other diseases at a certain period of life depends upon the cooperation 
of the generative organs, but while in the disposition to mammary cancer 
the internal secretion of the ovary is the main factor and suspension of 
breeding only subordinate, in the case of resistance to other death producing 
conditions the suspension of breeding is the main factor and the elimination 
of the ovarian function is only a subsidiary factor which merely acts through 
the suspension of breeding which it calls forth. The differences in the 
tendency to die from other causes than cancer which we find normally 
between different strains of mice are eliminated in mice which are prevented 
from breeding. All those strains in which breeding is prevented become 
approximately equally long lived. 


INHERITANCE OF CANCER IN MICE 


185 


11. As we stated more than 14 years ago, the endemic occurrence of cancer 
among animals is due to this hereditary transmission of the disposition to 
cancer. Infection with certain metazoon parasites which stimulate the 
tissues to proliferation may also play a part. The observations of Fibiger 
and others suggest this additional conclusion. 

12. While these statements apply directly only to animals, the evidence 
on hand makes it at least probable that in principle conditions are similar 
in man. Here also in all probability an intensity factor is hereditarily 
transmitted, which in many cases, however, has been equalized among 
different families as a result of long continued interbreeding. As to the 
increase in the cancerrate which seems to be so common an occurrence, we 
may suggest that as far as it is not due merely to improved diagnosis, it 
could be referred to a relatively greater frequency in the dominance of the 
parent with a tendency to a higher Tumorrate in the offspring. 


THE INHERITANCE OF A PREDISPOSITION TO CANCER 

IN MAN 


C. C. LITTLE 

Carnegie Institution of Washington, Cold Spring Harbor, Long Island, New York 

The work of Murray (1908), Haaland, (1908), Tyzzer (1909) and Slye 
(1913) have shown that a predisposition to the formation of neoplasms 
is inherited in mice. The exact type of inheritance is still a matter of debate 
although the weight of evidence points in the direction of a complicated 
Mendelian inheritance of many factors as the underlying causes. From 
the outset it should be made clear that inheritance in the direct sense of 
transmission is not involved. It is rather that certain individuals inherit a 
tendency towards unbalanced growth, following irritation from either 
internal or external causes, of various parts of the body. That such a pre¬ 
disposition is inherited in mice is beyond question. 

In the case of man there has been from the outset a certain amount of 
confusion whenever the question of the inheritance of cancer has been dis¬ 
cussed. This is largely due to the fact that a direct type of inheritance or 
transmission was sought for by some investigators while an effort to prove 
the inheritance of a tendency or predisposition to cancer was the goal of 
others. There seems to be little doubt that direct transmission of cancer is 
out of the question. No positive evidence for it can be said to exist. With 
the matter of a predisposition, however, the case is much more difficult to 
prove or to disprove. The result has been that pedigrees and statistical 
data have been advanced as evidence both for and against inheritance and 
we find well known medical investigators on both sides of the question. 

The present investigation aims to analyze statistical data obtained in the 
family history records of the Eugenics Record Office, Department of Genet¬ 
ics, Carnegie Institution of Washington, and it may be well at the outset to 
bring out certain points of advantage which it is believed these data possess. 

1. They are collected and recorded by that member or those members of 
any given family whose prime interest is the accurate recording of family 
traits, both physical and mental. This should be contrasted with the ordi¬ 
nary method of collecting statistics (insurance companies) where a particular 
individual may or may not be well informed concerning the history of his 

186 


INHERITANCE OF PREDISPOSITION TO CANCER 


187 


family. That is to say, ignorant, illiterate individuals may quite as often 
be the source of data as are critical and well informed ones. There is, in this 
case, no previous selection as there is bound to be in the case of the Eugenics 
Record Office blanks. 

2. There is no reason why the data in the Eugenics Record Office blanks 
should attempt to conceal the incidence of such a biological condition as the 
occurrence of cancer. Nor is there any reason why an individual should 
seek to exaggerate its occurrence. This is not true of such statistics as might 
be collected by Insurance Companies from applicants for policies. It is 
distinctly to the advantage of an applicant to minimize the occurrence of 
ailments among his or her ancestors. 

3. The information in the Eugenics Record Office blanks is far more 
extensive; covering a greater family tree than the best insurance company 
data can possibly be. 

4. The opportunity is given to study the incidence of cancer in any genera¬ 
tion or group of a family. This is essential for, as I have some years ago 
pointed out, it is quite possible to have true inheritance of a tendency to 
cancer and yet little or no available evidence from a statistical point of view 
if only the relationship of parent to offspring is considered. In other words 
it is necessary to have available information concerning the sibs of cancerous 
individuals to supplement and extend the results of analyzing the relation of 
parent to offspring. In this respect the Eugenics Record Office data are 
excellent. There are undoubtedly other advantages but those listed above 
appear to be sufficient to justify an analysis of the Eugenics Record Office 
data in an attempt to obtain evidence bearing on the inheritance of a pre¬ 
disposition to cancer. 

At this point it should be noted clearly that the term cancer is used to 
include all neoplasmic growths which have caused death. In other words, 
sarcomas and carcinomas are lumped together, and no effort is made to 
differentiate between them. This usage is admittedly a popular rather than 
an exact one, but since all data based on hearsay evidence rather than actual 
detailed autopsies must be somewhat indefinite in their very nature, a more 
accurate usage would be equivalent to the use in mathematics of a third or 
fourth decimal place, when only two were significant. 

In order to determine whether or not a hereditary tendency exists, it is 
necessary to establish an adequate control. In the case of this work, two 
sets of controls have been used. 

a. The table given by Hoffman (Mortality from Cancer throughout the 
World on the number of deaths from cancer per 100,000 inhabitants in the 


188 


C. C. LITTLE 


United States Registration Area (1903-1912) has been used. This table is 
divided by sex and age groups as follows: 


AGE AT DEATH 

HALES 

FEMALES 

Deaths from cancer 

Rate per 100,000 
population 

Deaths from cancer 

Rate per 100,000 
population 

Under 10 

1,170 

2.5 

984 

2.2 

10-24 

2,028 

3.1 

1,844 

2.8 

25-34 

3,757 

9.0 

7,891 

20.6 

35-44 

10,750 

32.3 

26,779 

89.0 

45-54 

24,431 

105.4 

46,669 

222.9 

55-64 

35,327 

257.4 

50,393 

386.4 

65-74 

33,745 

452.8 

43,010 

565.7 

75 and over 

18,381 

620.2 

24,601 

734.1 


From it the chance of any individual to die of “ cancer” can be directly 
calculated. In our tabulation, individuals still alive have all been recorded 
as negative, and have been given their individual values according to their 
respective age groups and their sex. They represent unfulfilled chances for 
the production of cancer. In addition the deaths from causes other than 
“ cancer” have been added, thus increasing greatly the difficulty of demon¬ 
strating an hereditary influence. 

The first group to be considered is composed of fraternities whose father 
was cancerous and whose mother was normal. 



TOTAL NON- 
CANCEROUS 

TOTAL 

CANCEROUS 

CANCEROUS 

INDIVIDUALS 

EXPECTED 

EXCESS OF 
OBSERVED 
OVER 
EXPECTED 

ODDS AGAINST EXCESS BEING 
DUE TO CHANCE ALONE 

Males. 

939 

778 

14 =*= 2.50 
10=*= 2.12 

0.85 

1.05 

13.15 

8.95 

More than 1350:1 
215:1 

Females. 


Total. 

1717 

24=*= 3.27 

1.90 

22.10 

More than 100,000:1 



It will be seen that in each of the three cases, the excess of cancerous in¬ 
dividuals among the progeny of cancerous fathers and normal mothers is 
striking. 

A similar result is observed when a tabulation of the immediate progeny of 
cancerous mothers and non-cancerous fathers is made. 





























INHERITANCE OF PREDISPOSITION TO CANCER 


189 



TOTAL NON- 
CANCEROUS 

TOTAL 

CANCEROUS 

CANCEROUS 

INDIVIDUALS 

EXPECTED 

EXCESS OF 
OBSERVED 
OVER 
EXPECTED 

ODDS AGAINST EXCESS BEING 
DUE TO CHANCE ALONE 

Males. 

553 

477 

12 =*= 2.33 
27=*= 3.40 

1.19 

1.59 

10.81 

25.41 

520:1 

Over 500,000:1 

Females. 


Total. 

1030 

39 =*= 4.13 

2.78 

36.22 

Over 1,500,000:1 



The final tabulation to be considered is that of fraternities in which at 
least one cancerous individual appears. If chance alone is operative, there 
is no reason why the sibs of such cancerous individuals should be cancerous 
any more frequently than is the population at large. Actually, however, a 
great excess is obtained. 



TOTAL NON- 
CANCEROUS 

TOTAL 

CANCEROUS 

CANCEROUS 

INDIVIDUALS 

EXPECTED 

EXCESS OF 
OBSERVED 
OVER 
EXPECTED 

ODDS AGAINST EXCESS BEING 
DUE TO CHANCE ALONE 

Total. 

2016 

48=*= 4.61 

5.80 

42.20 

Far over 1,500,000:1 



From the above three lines of evidence we may conclude that there exist 
in man one or more hereditary tendencies to the formation of malignant neo¬ 
plasms. 

b. The sibs of cancerous individuals can be contrasted with the general 
population as shown in the Eugenics Record Office data themselves as 
follows: 



GENERAL POPULATION 

SIBS OF CANCEROUS 

Non-cancerous. 

52,499 

657 

2016 

Cancerous. 

48 


Total. 

53,156 

1.23=*= .03 

2064 

Per cent cancerous. 

2.32=*= .22 



In addition, the progeny of one cancerous parent can be contrasted with 
the general population from which the families containing the cancerous 
parent in question has been subtracted. When this is done, the following 
result is obtained: 


General population 


Non-cancerous. 44,593 

Cancerous. 274 

Per cent cancerous.0.61=*= .02 



















































190 


C. C. LITTLE 


Immediate progeny of one cancerous parent 


Non-cancerous_ 

Cancerous. 

Per cent cancerous 


. 2,747 

. 63 

2.24 =*= 0.19 


Difference 1.591 =*= 0.191 (8.5 times the probable error.) 

From both these sources of data it may be concluded that a tendency to 
the formation of cancer is clearly inherited. The influence of inheritance is 
shown by the occurrence of a marked excess of “ cancerous” individuals over 
the rate of the general population, in (1) the sibs of “cancerous” individuals, 
(2) the progeny of “cancerous” mothers by “non-cancerous” fathers; (3) the 
progeny of “non-cancerous” mothers by “cancerous” fathers. 

The fact of inheritance is clear, but the type of inheritance needs further 
investigation. It does not appear to be simple Mendelian inheritance. 
This does not however, preclude the possibility that it will be found to be 
dependent upon multiple Mendelizing factors. 

The above findings have little or no direct bearing on the therapy of cancer. 
They should be simply taken to mean two things. (1) The case of man is 
closely parallel to that of the laboratory mammals already referred to. 
(2) If, therefore, the future brings out facts of general biological signifi¬ 
cance and therapeutic value concerning cancer in laboratory mammals, 
those facts bid fair, with a small amount of change, to be applicable to 
humans in at least some degree. Especial attention should therefore 
be given to the biological investigation of the cancer problem, as a series 
of growth phenomena, in laboratory mammals. The somewhat tedious 
question which one frequently hears as to the value of such investigations 
from a human viewpoint should be, in a large measure unnecessary. 





INHERITANCE OF EYE DEFECTS 


LUCIEN HOWE 
Buffalo , New York 

It is only possible here to call attention to a few of the more salient points 
concerning the subject. 

The first is the fact that effects of inheritance are shown in the eye. 
Everyone recognizes this in its color, position and outward characteristics. 
But it is also shown in the anatomy and physiology in so many ways and to 
such a degree as to make the eye practically a pathological museum of 
inheritance. A table of these inherited characteristics is shown here. A 
glance at the list shows how many and how varied they are. Anyone who 
may care to know more of the subject, is referred to a bibliography of heredi¬ 
tary eye defects, compiled by the writer and published a few months ago as 
bulletin 21 of the Eugenics Record Office of the Carnegie Institution of 
Washington. 

In passing it is worth while to observe that in the curvature of the cornea 
we have an opportunity to obtain, as perhaps nowhere else, an expression in 
figures of the degree of exactness of heredity. This is shown in astigmatism. 
It is well known that this defect consists in a difference in the curvature of 
the cornea in one meridian as compared with the other. In the ordinary 
form of astigmatism, the eye is not a globe but is shaped like a rubber ball 
which has been pressed from above down, or from side to side or at some 
oblique angle. In other words the radius of curvature in one direction is 
greater than in the other. Now, we can find the length of each radius by 
simply placing a suitable lens before the eye when testing the vision. The 
difference between these two radii may be, for example, not more than a 
few hundredths of a millimeter and yet that defect is sometimes transmitted 
from parents to offspring. Not only that, but the relative position of these 
two axes in the child is sometimes similar to that in the parent Other 
examples almost as marked as this, could be given if the limits of this paper 
permitted. The point is that heredity does show itself in many ways of 
rather unusual interest. The second point to which I would call attention 
is that the medical profession has until recently contented itself with simply 
recording the existence of this or that defect in certain families. Some of 

191 


192 


LUCIEN HOWE 


these histories, especially those given by Nettleship have been worked out 
with great care and detail. But during the last half dozen years, ophthal¬ 
mologists have awakened sufficiently to ask, what shall we consider 
heredity? Indeed, the terms “filial” and “congenital” and “hereditary” 
have been so confused as to mean little or nothing. The general conclusion 
is that the word “filial” should be dropped entirely, because so indefinite. 
That “congenital” should be used only in its true sense as a trait apparent 
at birth. So it remains for us to determine the meaning of the word heredi¬ 
tary. By that term we understand ordinarily the appearance of a certain 
characteristic, occurring more or less frequently in one generation after the 
other, following more or less closely the Mendelian sequence. But in 
studying these family histories, one especially has appeared, (and others 
seem to approach it) in which the number of those affected is so large as to 
be at variance with all of our former ideas concerning what we have called 
inheritance. This unusual family was reported by Dr. D. G. Risley, in the 
Journal of the American Medical Association for April 17,1915. The defect 
consisted in the absence of part of the iris (coloboma iridis) or, in some, entire 
absence of that membrane. The unusually large proportion of the defec¬ 
tives was contrary to the law of Mendel or of any one else. The only ex¬ 
planation which can be offered is that the defect may be one of those rather 
obscure infections which is transmitted in this case by placental circulation 
or through the germ itself from generation to generation. Naturally one 
thinks first of a syphilitic infection. There seems no possible reason why 
that disease should produce a partial or entire absence of the iris—but in¬ 
vestigations are now started concerning this point. Since Parodi in 1907 
showed that rabbits can be inoculated with the Treponema pallidum, 
occasional attempts have been made to obtain the eye lesions and then, by 
breeding, observe the eye changes, if any, in subsequent generations, but a 
rabbit is here shown which after inoculation, has lost both eyes, and may yet 
become the sire of other syphilitic rabbits, possibly with partial or total 
absence of the irides. 

But the special object of this paper is, to call attention to a series of ex¬ 
periments which have been made known this year by Professor Michael F. 
Guyer of the University of Winconsin. A short account of these was 
published in the Journal of Experimental Zoology August 20 of last year, 
and mention made of them elsewhere. 

The statements were in brief that it was possible by serologic methods to 
so influence the blood supply of a pregnant rabbit that a part of the litter at 
least, would be born with imperfect eyes, and that similar imperfections 
could be transmitted from generation to generation. The novelty and im- 


INHERITANCE OF EYE DEFECTS 


193 


portance of that was evident. But in the published statements some de¬ 
tails of the process were not described, and the pathological conditions 
thus transmitted described only as “imperfections,” “cataract” and “de¬ 
generation.” It was with much pleasure therefore that I embraced an 
opportunity to go to Madison in order to see these pathological conditions 
as they appeared when viewed with the ophthalmoscope, in a considerable 
number of these rabbits. The subject proved so interesting and Professor 
Guyer was so kind in furnishing me with parent stock that I am continuing 
the breeding experiments. 

In order to understand the principles involved it is desirable to recall 
briefly the reasons and method of the procedure. It has long been known 
that extracts of certain organs of the body of one animal (A) could be in¬ 
jected into a second animal (B) to a point which we might call saturation or 
technically, until the second animal was sensitized. Then if the serum of 
(B) is injected into another specimen of the same kind as (A), that serum 
from (B) would exercise a certain specific influence upon the organ corre¬ 
sponding to that from which the extract or emulsion was made of the animal 
(A). This of course is a familiar procedure in serology. 

Now the procedure of Guyer was to kill three or four rabbits by placing 
them in a gas chamber, to extract the lenses, make an emulsion of the lenses 
and inject that emulsion into a fowl up to the point of full sensitization. 

The next step was to inject the serum of that sensitized blood of the fowl 
into a female rabbit at the time when it would be most apt to effect the eyes 
of her offspring. Now we know that the point in the head of the foetus 
which is to become the eye, receives its greatest supply of blood about the 
twelfth to the fifteenth day after the mating. Therefore after bleeding the 
sensitized fowl, Guyer injected its serum into the veins of the ear or into a 
rabbit which he had reason to suppose was pregnant about twelve to fifteen 
days. 

After numerous failures some of the offspring showed decided defects of 
the eye. Three of these rabbits given to me by Professor Guyer are brought 
from my small collection in Buffalo for demonstration here. Described 
in general terms, one has one eye cataractous, the other partly atrophied. 
The second is practically in a similar condition. The third has only the 
stump of each eye remaining. They were all born in this condition. The 
accompanying charts show the lines of heredity in two families. Each in¬ 
dividual is numbered. 

In order to make sure that the blindness is artificially produced and not 
the result of breeding from some accidental defect, care was. taken with one 
of these families to obtain original breeding stock from another state. For 


194 


LUCIEN HOWE 


when a male with defective eyes is bred to a normal female, the first litter is 
nearly or entirely normal. But when females of that litter are bred again, 
to a defective-eyed father, those females produce some defective-eyed off¬ 
spring. Evidently there is no placental relation between those grand¬ 
children and the defective-eyed grandparents. The daughters of the first 
generation are merely carriers of the hereditary defect. This is entirely 
analogous to what we find so constantly in color blindness and some other 
defects in eyes of the human species. In a short paper of this kind it is 
impossible and also unnecessary to describe in detail the several structural 
defects of the eye which seem to be hereditary, especially those visible with 
the ophthalmoscope, those being interesting only to ophthalmologists. 

It is proper however, to observe this selective action of the lens or what 
Hektoen of Chicago has recently described as the specific precipitin reac¬ 
tion of the lens, is an important factor in these evidences of heredity. More¬ 
over they seem to furnish some corroborative testimony concerning Elsch- 
nig’s theory of the serological nature of sympathetic ophthalmia. Finally 
if we could learn the mechanism of the destructive processes it is quite 
possible that by reversing the method we may obtain a hint at least as to 
improved forms of treatment of this or other diseases of the eye. 


NEW DATA ON THE GENESIS OF TWINS 
R. A. FISHER 

Rothemsted Experimental Station, HarPenden, England 

The subject of the genesis of human twins, apart from its intrinsic interest 
has a special importance for eugenists. According to the commonly ac¬ 
cepted doctrine twins are of two kinds: the so called uniovular or identical 
twins, which are supposed to have the whole of their heritable nature in 
common, and the so called biovular or fraternal twins which are supposed 
to be related genetically no more closely than ordinary brothers and sisters. 
It is clear that, if this view be correct, human twins provide a test case for 
measuring the importance of hereditary factors in the determination of 
human characters. For by measuring the degree of resemblance in any 
character between identical twins, we might obtain an upper limit for the 
effects of heredity, while by comparing the resemblance of fraternal twins, 
with that known to exist between ordinary brothers and sisters, we might 
measure the effects of early environment and especially of intrauterine 
environment. The importance of human twins as a test case for eugenics 
was appreciated by Galton, and since the publication of his work, the con¬ 
clusions at which he arrived have been repeatedly quoted by other writers. 

Current opinion generally accepts the division of twins into the two classes 
distinguished above, classes both of which have analogues in the animal 
kingdom. Great disparity of opinion exists, however, as to the frequency 
with which these two classes occur. Some biologists believe that identical 
twins are relatively frequent. Extensive statistics show that twins of 
unlike sex occur in about 3 cases out of 8; and if we argue that of the 
fraternal twins nearly 50 per cent must be of unlike sex while all identical 
twins must be of like sex, we are led to the conclusion that about one pair 
of twins in 4, must be an identical pair. Others, however, ignoring the 
distribution of sex, believe that identical twins are excessively rare. It is 
generally believed that the two types of twins may be distinguished by an 
examination of the fetal membranes, but the proportions arrived at in this 
way by gynecologists are excessively divergent, and according to Daven¬ 
port, twins of unlike sex are frequently reported to be uniovular in con¬ 
tradiction of the current theory. 


195 


196 


R. A. FISHER 


The only extensive body of measurements of twins was taken by Thorn¬ 
dike, and published sixteen years ago. It is not sufficiently recognized 
that his results are clearly inconsistent with both of the views currently 
held. The 50 pairs of twins observed by Thorndike do not fall into two 
distinct groups, but show themselves on examination to be a single homo¬ 
geneous group. The most delicate tests prove that those pairs of twins 
most like in any one trait, are not more alike than others in other traits. 
The whole group arises apparently from a single mode of genesis; and those 
of unlike sex are not sensibly less alike than those of like sex. If we adhere 
to the theory of the dual origin of human twins, we must conclude from 
Thorndike’s data either, that (1) all or nearly all the twins examined 
were “fraternal,” in spite of the facts (a) that the proportion of like sex is 
much greater than 0.5, and ( b ) that the level of resemblance in all traits 
is much higher than that found between brothers and sisters, or (2) that all 
or nearly all were “identical” in spite of the facts {a) that 9 pairs were of 
unlike sex, and ( b ) that the general level of resemblance is considerably less 
than we should, for other reasons, expect between genetically identical 
individuals; for example, several pairs differ considerably in eye color and 
hair color. 

For these reasons, after a statistical examination of Thorndike’s data, 
I have brought forward (Genetics, 4, 489, 499, September, 1919) the 
tentative suggestion that the ordinary mode of genesis of human twins is 
intermediate in character between the fraternal and identical modes of 
genesis, and in particular that while the maternal inheritance is identical, 
the twins being derived from a common ovum, the paternal inheritance is 
merely fraternal, the twins being derived from different spermatozoa. 

The purpose of the present note is to present such new data as I have 
been able to collect in testing this particular suggestion. 

Children of the same parents may differ in a Mendelian factor, if either 
or both of the parents are heterozygous; these differences will be apparent 
if neither parent is a homozygous dominant. To make clear how the 
suggestion may be tested, the following table shows the parentage of unlike 
twins produced, on each of the possible modes of genesis, according as the 
mother, the father, or both parents are heterozygous. 



MOTHER 

HETEROZYGOUS 

FATHER 

HETEROZYGOUS 

BOTH PARENTS 
HETEROZYGOUS 

Fraternal. 

50 

50 

37.5 

Semi-identical. 

0 

50 

25.0 

Identical. 

0 

0 

0 













NEW DATA ON THE GENESIS OF TWINS 


197 


It is apparent from this table that if twins are readily produced in the 
manner here tentatively suggested, we may lay down the rule, which 
should be capable of definite proof or disproof, that all twins unlike in any 
Mendelian factor, have fathers heterozygous in that factor. 

The most suitable factor to choose for carrying out this test would seem 
to be eye color; for although recent observations have thrown doubt upon 
the simplicity of the inheritance of yellow pigment on the iris, it appears in 
the majority of stocks to be inherited as a simple dominant. Valuable 
cases also occur occasionally in some rare defects, which are known 
to be due to simple Mendelian factors; of these a number of cases are on 
record in which albinism appears in one twin and not in the other. The 
cases which I have hitherto obtained in which twins have been observed to 
differ in what is believed to be a simple Mendelian factor, are set out below: 


Albinism... 
Eye color.. 
Other cases 


Total. 


FATHER 

HETEROZYGOUS 

FATHER 

HOMOZYGOUS 

4 

0 

2 

0 

2 

0 

8 

0 


It will be seen that no case has as yet come to light in which the father is 
homozygous; cases in which both parents are heterozygous have not here 
been recorded, so that on the current theory we should expect to find the 
father homozygous in one-half of the cases. 

The above numbers are far from conclusive, and have been put forward 
to show that the subject should be thoroughly investigated. We hope 
that this may be done in London, where over a thousand pairs of twdns 
could rapidly be obtained; but at present the work is delayed for lack of 
funds. In the second place, it should be possible quickly to ascertain 
whether or not the father of unlike twins is always heterozygous. One 
hundred accurately recorded cases should settle the whole question; of 
course out of a hundred cases we must expect a certain number of excep¬ 
tions owing to wrongly ascribed paternity, but in such a number it would be 
amply clear, whether or no there were a real difference between the paternal 
and the maternal inheritance of twins. 















MAIN RESULTS OF A STATISTICAL INVESTIGATION OF 
FINGER PRINTS FROM 24,518 INDIVIDUALS 

KRISTINE BONNEVIE 
Institute of Genetics , Kristiania 

As our knowledge of human inheritance proceeds it will on various 
points be taken into the service also of the court of justice, especially in 
questions about paternity. The cases are, however, as yet very few in 
which a decisive proof of paternity could be afforded through investigation 
of hereditary characters of persons in question (see Mohr 1921). 

The number of requests from lawyers for a hereditary investigation of 
their clients is nevertheless rapidly growing, and therefore it would be of 
importance for experts to know the law of inheritance of a series of external 
human characters, as a basis of their decision of whether or not these charac¬ 
ters might be used for a statement of relationship between two persons. 

It is but natural that a character like that of the “finger prints” 
which has already performed so important services in questions of identifica¬ 
tion, should come in the first row also in an investigation of inheritable 
characters. And there exist indeed, already several indications of the 
heredity of general types of patterns on both hands and feet, even if the 
details of these patterns always have been shown to be characteristic of a 
single person only. 

Our modern knowledge about the patterns formed by papillary ridges 
on the fingers of man is founded above all upon the results of Galton laid 
down in a series of papers among which his book on “Finger Prints,” 1892, 
should be mentioned in the first row. A voluminous literature, contribu¬ 
tions to which have been given from many different countries, has followed 
the publications of Galton. Of special interest are the papers by Whipple 
(1904), by Schlaginhaufen (1905-1906) and by Wilder (1916), the two 
last named papers containing also full lists of the literature upon this 
subject. 

With regard to the heredity of papillary patterns Galton has in his famous 
book on “Finger Prints” (1892) already treated the question in a chapter 
containing a very valuable material. Through a statistical treatment of 
the finger patterns of 150 fraternal couplets, further through an investiga- 

198 


STATISTICAL INVESTIGATION OF FINGER PRINTS 


199 


tion of 17 sets of twins and finally also through a comparison of children with 
their parents in cases in which both parents were “like patterned”—Gal ton 
has reached a conclusion expressed in the following words (p. 189): “The 
decided tendency to hereditary transmission cannot be gainsaid in the face 
of these results, but the number of cases is too few to justify quantitative 
conclusions.” 

More conclusive evidence is given by Wilder (1916, 1919), although his 
attention has been directed more on palm and sole patterns than on those of 
the fingers. 

His results are reached through an investigation partly upon the hands 
and feet of twins, partly also upon whole families. 

The papillary patterns of the fingers have, however, not yet been subject 
to special investigations with regard to their heredity. And before this 
question can be definitely solved it will be necessary to settle a series of other 
questions with regard to the statistical occurrence of the different types of 
patterns upon each of the fingers, to the correspondence between different 
fingers and also to the symmetry between the fingers of the right and the 
left hands with regard to their types of pattern. 

Without knowledge about these facts one will have no real base for a 
sound criticism of the results of family investigations. 

Galton (1892) has, upon this point also, made the beginning through a 
universal statistical treatment of the finger-patterns of 500 individuals. 
His results which on many points seem extraordinary but which have in 
the main lines been stated through my investigation will be treated together 
with the results of the latter. 

The statistics of this paper (tables 1 and 2) are based upon the material 
of The Office of Identification of Kristiania, which by its Director was very 
kindly offered at my disposal. The finger prints of this office are taken from 
24,518 individuals and counting, therefore 10 times this number of fingers. 
It is classified according to the scheme generally adopted for an identifica¬ 
tion of criminals (see Henry 1901), according to the type of pattern on each 
finger on both hands. 

Three types of patterns are known as whorls , loops and arches. 

The whorls are characterized through the existence of two deltas , that is 
on each side of the pattern one ridge dichotomously divided forming a 
triradius so as to embrace the central part of the pattern, in which the ridges 
may be circular, forming a single or a double spiral or even other more 
irregular figures. 

In the loops there is only one delta while one or more ridges are forming 
loops opening towards the side of the finger opposite to that of the delta. 


200 


KRISTINE BONNEVIE 


The loops may open towards the radial or towards the ulnar side of each 
finger and according to this they will in this paper be called radial or ulnar 
loops. 

The arches represent the simplest patterns, the ridges here running in 
more or less deep bows from side to side of the finger without making any 
backward turn or twist. There is no real delta in the arch. 

As already mentioned, the statistics of this paper are based upon 24,518 
individuals, or a number of 245,180 fingers. The formulae of all individuals 
have been dissolved so as to give the type of every single finger and after 
this the number of fingers with one and the same type has been counted for 
application and combination in different ways. 

It is seen from table 1 that the three types of finger-patterns are very 
unlike with regard to their numerical appearance within the whole material 
(245,180 fingers), the whorls (62,883) appearing in a percentage of 25.65, the 
loops (164,150) in 66.95 per cent, while the percentage of arches (18,147) 
is only 7.4. Such a difference proved in a material large enough to exclude 
every influence of chance, at once indicates that the finger-patterns in their 
occurrence are under the control of some constitutional law. 

In table 1 we find also a conspicuous difference in the occurrence of 
radial and ulnar loops, the latter appearing in a percentage of 61.14 of the 
number of fingers, while the percentage of radial loops is only 5.81. 

The distribution of pattern-types between right and left hands also 
proves to be peculiar, the whorls being considerably more numerous on 
right hands (29.38 per cent against 21.92 per cent on left hands), while 
arches (6.92 per cent to 7.88 per cent) and especially also loops (63.70 
per cent to 70.2 per cent) are more richly represented on left hands. This 
difference is, within the loops, due to the ulnar loops only (57.76 per cent to 
64.52 per cent) while the occurrence of radial loops (5.94 per cent to 5.68 
per cent) is practically the same on both sides. 

The above statistics refer to the whole material of fingers taken together. 
The picture here given is very much deepened through a consideration also 
of each finger separately. 

It is from the figures of each horizontal line (table 1) evident that digits 
I-V all have their special characteristics with regard to the statistical 
Occurrence of the pattern-types. 

Very obvious is the general similarity between digits I and IV , both 
having a very high percentage of whorls especially upon right hands, and 
both having also a very low number of radial loops , while ulnar loops and 
arches also show values considerably below the average especially on right 
hands. 


TABLE 


STATISTICAL INVESTIGATION OF FINGER PRINTS 


201 
















































































202 


KRISTINE BONNEVIE 


Digit II gives a picture very different from the one here mentioned. 
Whorls are occurring in a percentage somewhat above the average but very 
equally distributed on right and left hands; this approximate statistical 
correspondence between digit II of right and left hands is valid also for the 
other patterns. 

But the most striking feature in the characteristics of digit II is that of the 
exceedingly high percentage of radial loops and also of arches , both pattern- 
types somewhat, but not very much, more numerous on right hands. The 
ulnar loops , in return, are occurring in a percentage far below the average. 

Digit III shows a relatively low number of whorls and a high one of ulnar 
loops , while radial loops give values below the average for this pattern. 
Arches appear, also here, remarkably frequently even if their values are 
considerably below those of digit II. 

Digit V finally, has high numbers of ulnar loops only, and low percentages 
of all other patterns. But even here some peculiarities of their distribution 
should be noted, namely the very considerable surplus of whorls upon right 
hands as compared with the left ones,—and also the peculiar occurrence of 
radial loops. Their percentage of occurrence is below the average although 
higher than that of digits I and IV, but their distribution upon right and left 
hands is very unlike radial loops scarcely occurring on digit V of right 
hands but in a percentage higher than all other fingers except digit II 
on the left hands. 

The characteristics of the various fingers as now studied on table 1 re¬ 
ceives an interesting supplement through a look also at table 2, showing 
the statistical distribution of each pattern type; their occurrence on the 
fingers is here given in percentage of the absolute number of the type contained 
in the whole material. The average number upon each of the 10 fingers 
would then, of course, be exactly 10 per cent of each type. 

The whorls are distributed with high ratios upon digits I and IV of 
right hands and with low ones upon digits III and V of both hands, while 
upon the fingers not here mentioned their occurrence is little above the 
average. 

The radial loops are very particular in their distribution, more than four- 
fifths of all of them being found on digit II, the right hand with the highest 
percentage. On all other, fingers radial loops are very rare, the lowest 
percentage being found on digit V of right hands, while the same finger 
of left hands has a number of radial loops higher than that of digits 
I, III and IV. 

Ulnar loops show, in return, upon digit II an occurrence far below the 
average, while on digits III and V it is rising above this point; upon digits 
I and IV the percentages of ulnar loops give very near average values. 


TABLE 


STATISTICAL INVESTIGATION OE FINGER PRINTS 


203 



o o 
o o 

o o 
o o 
r* 

o o 
o o 

o o 
o o 

T- T“ 

o o 

o O 

o o 

O O 
■v* ■*- 

O o 
o o 

o o 
o o 

r- T- 

o o 
o o 

o o 
o o 

t—■ r— 


3 

f 5 
• s 

f 

Vi\ 

•4- 

f'i 

<o 

«> 

^v> 

Vi 

to 

tO 

in 

... 


«3 

O 

4~' 

°° 

J 

*o 

s' 

|, 

♦o 

«o 

v> 

v> 

*-G) 

1 

V* 

«o 

'~o 

cr 

c-C 

o' 

I 

v> 

V> 

r- 

*~i 

v- 

/ 

r-’ 

C\| 

T— 

o 

N 

r- 

1 

N 

o 

V>| 

* 

p 

4f 

* 

Q 

f° 

f 

A 

<r* 

«o 

t— 

«*> 

cr- 

\ 

N 

.3 

o' 

o 

r* 

O 

cr 

o' 
\— 

o 

rvf 

J 

o 

o 

T~ 

N 

~T 

V) 

o 

v> 

- 

<o 

<^4 

a-' 


X* 

n 

f 

o 

V> 

r-* 

•t- 

Vo’ 

<*> 

O 

/ 


o 

<o 

Vi 

cr- 

x— 

3 

t~ 

r 

/ 

v> 

tK 

1 

.t 

r 

& 

i< 

cO 

§■ 

•* 

Q 

<3 

4* 

\ 

4- 

T— 

3-' 

>D 

cr 

uS 

•4 

r~' 

'-n 

o 

r- 

■T— 

\ 

O 

vS 

T- 

cr 1 

Osf 

f> 

«JO 

Vi 

O 

7^4 

V* 

«v 

s*5 

«o 

. 

r- 

O 

>X 

T-* 

l 

O 

cv 

rJ 

CT 

O 

‘■v 

J 

°/ 

Vi] 

wV] 

4 

b 

& 

i. 

c\7 

S' 

<y 

o 

5^ 

</■> 

h 

J 

<N 

«3 

«r> 

f>» 

l* 

A 

'■O 

’•n 

«o 

° 

J 

no 

i 

L 

CO 

•0 


o 

'■o 

4- 

-*A 

o 

•T 

\P 

r- 

Vj 

fNi 

o 

Cv< 

4- 

4 

j 

°o 

o 

V> 

*4-* 

J 

f ° 

T- 

c- 

r-' 

(3 s - 

co" 

J 

£ 

r 

4^ 

f 

4 

oo 

O 

v* 

1 ^ 

cO 

o 

'■o 

o' 

Y“ 

o 

v) 

o' 

T~ 

p- 

<N? 

cr- 

cr-' 

* 

cnC 

J 

o 

r- 

OO 

15 . 

3 —* 

f 

crs 

4- 

«N 

>3 

V 

} 

r»-* 

<Nj 

O' 

l 

T"*" 

O 

0“ 

j 

t cr 

V 

«o 

oo 

<o 

!o 

^o' 

j 

o 

V 

r 

a- 


«o 

r- 

«r^ 

£ 

\ 

cr 

<aO 

,g 

A 

r- 

‘jo 

th 

w 

r-' 

*• 

^ 

r 

-O 

r-l 

lo 

r-' 

<o 

o 

T— 

J 

CN? 

CO 

*o 

r- 

r- 

-3 

=t* 

J 

. o 
o 

V- 

o 

T 

r- 

N 

r-~ 

V> 

> 

Jo 

^r* 

o 

T“ 

'-O 

! 

/ 

T* 


5 c 
,\ 

> 

■> 

■? 

3 

] 


r* O-A 

r > 

if J 

? 

°5 


c* ^ 

<vc Icko^ 













































































































204 


KRISTINE BONNEVIE 


The arches finally, like the radial loops, have their highest values on digit 
II, nearly one half of all arches occurring on this finger. But also on digit 
III we find arches richly represented, and then in sinking values upon digit 
I, IV and V. 

Many questions arise through an attentive consideration of the figures on 
tables 1 and 2. Above all, however, answers are wanted to the following 
questions: 

Which are the reasons of: 

1. The general statistic similarity between the first finger and the fourth? 

2. The great frequency of whorls upon the fingers just mentioned as well 
as the very striking surplus of their percentage upon the right hands, 
a feature which is visible on all fingers but especially valid also for 
digit V? 

3. The peculiarities of the second finger, viz. (a) the surprisingly high 
percentage of radial loops , with a considerable surplus upon right hands; 
and (b) the high value also of the percentage of arches. 

4. The relatively high percentage of arches and, less so, of radial loops 
upon the third finger? 

5. The strange difference between right and left hands with regard to 
radial loops upon digit V, their value on the left side being more than 18 
times that of the right. 

The answers of these questions can certainly not yet be given, and even a 
thorough discussion of them would lie outside the scope of this paper. A 
few words may be mentioned, however, with regard to the directions in 
which the answers might be sought. 

The existence of papillary patterns on the human palms and soles has, as 
before mentioned, in full agreement been explained phylogenetically as a 
remainder from patterns on the elevated pads of early ancestors, especially 
among prosimia and marsupialia (Klaatsch, 1887-1888; Whipple 1904; 
Schlaginhaufen, 1905; Wilder 1916). 

With regard to their function two different theories have been maintained, 
the one (Whipple) considering the papillary patterns as organs of a mechani¬ 
cal use (“friction-ridges”), the other theory attaching more importance to 
the sensory function of the ridges (“Figure tactiles” Schlaginhaufen), 
both theories agreeing, however, in the conclusion that the function of 
papillary patterns is best fulfilled, when the direction of their ridges is 
one vertical to the direction of pressure against the object to be touched. 

When trying to answer the questions above raised, one has then to con¬ 
sider the interaction between two different forces, the one conservative, 
preserving phylogenetically older conditions, the other force adaptive, 


STATISTICAL INVESTIGATION OE FINGER PRINTS 


205 


changing these conditions according to the function of each finger. A 
correct understanding of the effects of these two causes will be of decisive 
importance also to the whole question about heredity of finger-patterns. 

The peculiarities with regard to the statistical occurrence of pattern- 
types may be divided into two groups, those regarding the whorls upon 
digits I and IV (question 1-2) and on the other side those of the radial 
loops and arches upon digits II-III (question 3-4). The peculiarities of 
digit V will enter partly into the first and partly into the second of these 
groups. 

Considering the first group of questions, the similarity between digit 
I and digit IV and the very frequent occurrence of whorls especially on these 
fingers of right hands, it seems evident that the causes of these peculiarities 
should be sought before all in the phytogeny of patterns. A functional 
adaptation is, that is true, easily imagined for the first finger with its many- 
fold use, but an explanation of this kind will scarcely go for the fourth finger r 
which on the human hand, is perhaps less active than any of the other fingers. 
In this finger, therefore, phylogenetically old characters may especially 
have been preserved and if similar conditions are found in the first finger the 
same explanation should probably also be found valid for both fingers. 

With regard to the phylogeny of pattern-types Wilder (1916), basing on 
the investigations of Whipple (1904), maintains that the patterns of human 
hands represent (p. 235) “ every stage in the process of development, from a 
whorl of concentric lines with a definite number of embracing triradii, to a 
condition in which core and triradii are completely lost and the surface is 
covered by simple parallel ridges, straight or slightly curved.” And further 
also that “from comparative study of other primates it is plain that the 
course of evolution has been from the involved condition the whorl, which is 
distinctly simian, through every stage in the reduction of the pattern, to its 
final complete effacement.” 

If this be true, it is of great interest to find the whorl, as the phylogeneti¬ 
cally oldest pattern, so richly represented on the fourth finger. One must 
only ask why this peculiarity of the fourth finger should be found so nearly 
repeated in the patterns also of the first? 

Without having made any investigations upon this point myself, I can 
again only look at the statistic facts of my tables in light of the results al¬ 
ready reached by the investigators mentioned. All these authors agree 
in the consideration of the papillary patterns as reminiscences of elevated 
pads in the ancestors, the same elevated pads which are found also in human 
embryos and young children (Johnson, 1899, Retzius, 1904), and therefore 
also as indications (Whipple, 1904, p. 332) “that at the same time when 


206 


KRISTINE BONNEVIE 


ridges first developed in that region the pad had the elevated form of the 
typical walking pad.” Miss Whipple continues: “The second principle is 
that the relative constancy of occurrence of the typical pattern in a species 
or a group of related species, may be used as a criterion for determining 
the relative length of time during which various pads were retained in their 
typical form. ” 

The same principle being made valid also for the various fingers of one 
species the pads of human ancestors should he supposed to have existed longer 
upon the first and the fourth finger than upon any of the others. 

This is of interest in view of the fact that just among prosimians and 
marsupialias, the mammal groups which are supposed to be most nearly 
related to the row of human ancestors, many examples are found of a high 
development of the fourth finger, this finger being together with the first used 
by the climbing species for grasping round branches of trees. 

But there is also another feature of the statistics regarding whorls on 
digits I and IV which needs an explanation, viz., that of the great surplus of 
whorls on right hands. If the principle of Whipple, just mentioned, is 
applied also here, then the conclusion would be that the elevated pads of 
first and fourth fingers should have existed longer upon the right hands than 
upon the left ones. This conclusion should, in case, be made valid also for 
digit V , in which there is an equally great surplus of whorls on right hands. 

Personally I have no real base for taking a standpoint to the conclusions 
here drawn. They are logical consequences, however, of the theories 
maintained by Whipple as well as by Wilder and based upon the results 
also of Schlaginhaufen with regard to the phylogeny of finger-patterns. 

A difference between the first finger and the fourth with regard to the 
statistical appearance of whorls, these being more numerous (44.98 per 
cent of the fingers) upon digit IV of right hands than upon digit I (41.66 
per cent), may be an expression only of the more conservative conditions of 
the fourth finger. 

The second group of questions, regarding the peculiarities of digits II and 
III, contains problems of a very different nature. Here we must look for 
the answers not so much in the phylogeny of patterns as rather in the 
physiology of these special fingers. No other finger of the human hand is 
of a use so varied and extensive as just digit II and also the third finger 
very often partakes of its task in a position opposed to digit I. The high 
percentage of radial loops and of arches on digits II and III, therefore, may 
a priori be looked upon as a phenomenon of adaptation, and also here the 
results of previous investigators are in a remarkable way supported by the 
statistical facts. 


STATISTICAL INVESTIGATION OF FINGER PRINTS 


207 


As already mentioned both existing theories upon the function of papil¬ 
lary ridges maintain that the direction of the ridges ought to cross the line 
of greatest pressure. And the tactile theory maintains further, based upon 
experiment that the sensory effect of two parallel ridges is greater when 
both belong to one and the same loop than when they are independent of 
each other (Fere, 1895; Schlaginhaufen, 1905). 

Remembering the position of the second finger when working alone in 
opposition to the first one, it seems evident that the radial side of digit II 
and its papillary pattern should be of great importance, whether the func¬ 
tion of those lines be of a mechanical or of a sensory nature. Among the 
different pattern-types, therefore, the ulnar loops will be the ones least 
useful, their ridges running away from the radial side of the finger. Whorls 
and arches , the peripheral ridges of which patterns are running out parallel 
on both sides of the finger, seem here to be of equal use as compared with each 
other. But no other pattern would for the special use of the second finger 
make better service than the radial loops , the ridges of the radial side of the 
finger here being combined into pairs as arms of one and the same loop. 

This consideration of the functional importance of different papillary 
patterns agrees exceedingly well with their statistical distribution upon the 
second finger. The number of ulnar loops is here very considerably lower 
than upon any other finger, making (table 1) only 30.66 per cent of the 
number of fingers while 58.71 per cent (digit IV) is the lowest value upon 
other fingers. As shown in table 2,4 to 5 per cent only of all ulnar loops 
are found on the second fingers of right and left hands instead of the average 
10 per cent. The whorls give values not far above the average while arches 
and especially also radial loops are represented in extraordinarily high num¬ 
bers, 44.5 per cent of all arches and 82.57 per cent of all radial loops being 
found upon second fingers only (table 2). 

Although the statistical difference of pattern-types between right and 
left hands is considerably less evident on the second finger than on digits I, 
IV and V, it is interesting to note that also here a difference exists, the 
functionally most effective patterns (radial loops and arches) showing a 
considerable surplus on right hands. A parallel to this phenomenon is 
shown by Wilder (1916) to exist with regard to the formula of the human 
palm, the functionally most advanced configuration being considerably more 
richly represented on right than on left hands. Wilder draws in the same 
paper the conclusion (p. 236) “that this result has been gained since the 
adaption of right-handedness. ” 

What has been said here about digit II is valid, to a certain degree, also 
with regard to the third finger. Also here we find the finger engaging its 


208 


KRISTINE BONNEVIE 


radial side when working together with digit II, opposed to digit I, and also 
here we find the percentage of arches considerably above the average (11.03 
per cent to the average 7.4 per cent, table 2). The radial loops , however, so 
peculiar to the second finger, are here not strongly represented (2.31 per 
cent of the fingers), even if more so than on digits IV and V. 

A few words should be said also with regard to digit V, agreeing with the 
first named group of fingers (digits I and IV) in the great surplus of whorls 
on right hands (13.72 per cent of the fingers) as compared with the left 
hands (8.30 per cent),—and at the same time entering into the second 
group (digits II and III) on behalf of a peculiarity in the occurrence of 
radial loops. 

If the explanation given for the statistics of whorls upon digits I and IV 
shall prove to be correct, then the same reference to the phylogeny of 
finger-patterns will be valid also here. The great difference in the number 
of whorls on right and left side would, therefore, according to Whipple 
(1904) indicate that the inflation of the elevated pads developed more 
rapidly on the left side. Upon this side, therefore, the reduction of the 
original finger-pattern, the whorl, has set in earlier and the number of other 
patterns, loops and arches is higher on left hands (tables 1 to 3). It is at 
first glance surprising that the number of radial loops upon digit V (5.62 
per cent of all radial loops) should be so much higher than those of digit I 
(1.15 per cent) and digit IV (2.70 per cent). But it is, for example, by 
grasping round a ball, very easy to state that even if the use of digit V is upon 
the whole very restricted, its radial side is the one generally used. The 
peculiar difference between right and left hands with regard to the occur¬ 
rence of radial loops upon digit V, 0.29 per cent and 5.33 per cent respec¬ 
tively, stands as a fact the explanation of which should be sought through 
future investigations. 

The statistical facts now being analyzed, we may turn to the conclusions 
which will be of direct importance to a hereditary treatment of a material 
of finger patterns. 

Many cases may be demonstrated which at first glance seem to prove 
that heredity plays an important part in the distribution of pattern-types. 
A critical survey of these cases in light of the statistical results shows, 
however, that their value as proofs of heredity is very different. A few 
examples from the material at my disposal will illustrate this. 

Experiment 1. Two brothers are found with ulnar loops on all fingers of 
both hands. The pattern-formula of this example occurs in 4.29 per cent 
of all individuals, that is: Among each 50 persons, at least 2 should be 
expected to have this formula. The occurrence of two such brothers, 
therefore, cannot be considered as a proof of heredity. 


STATISTICAL INVESTIGATION OF FINGER PRINTS 


209 


Experiment 2. Two brothers have arches on all fingers of both hands. 
Arches on all fingers of both hands is a very rare occurrence found in no 
more than 0.18 per cent of individuals, that is: more than 1100 persons 
would be necessary to statistically secure the presence of 2 individuals with 
the formula mentioned. Its occurrence in two brothers, therefore, must be 
considered a very strong indication of heredity. 

Experiment 3. Three brothers have all a radial loop on the second finger 
of their right hands and ulnar loops on all other fingers. A radial loop on 
digit II of the right hand occurs in 25.73 per cent of individuals, ulnar 
loops on all fingers of one hand in 14.29 per cent. Even if I cannot tell the 
exact percentage of the occurrence of a formula like that of the example it 
is, therefore, quite sure that such a formula is not at all rare. Among a 
hundred persons we might easily happen to find three with finger patterns 
like those of the brothers mentioned, and they will, therefore, give no 
definite answer in questions about heredity. 

The examples here mentioned will suffice to show that hereditary investi¬ 
gations upon finger-patterns should comprehend first of all a critical survey 
of the material, basing on the statistical occurrence of each pattern upon 
different fingers. If in this complicated matter clear results shall ever be 
reached, the work ought to begin with patterns of a relatively rare occur¬ 
rence, as arches or radial loops. 

Besides the statistical data the influences of the phylogeny as well as of 
the physiology of finger-patterns should be considered in order to get a real 
understanding of the constitutional forces working in their heredity. 

Basing upon Whipple’s theory of the whorls as the typical pattern of 
elevated pads, a theory which, as mentioned above, has got a strong support 
in some of the statistical results, and of the other patterns as a series of 
reduction stages from the whorls, our results with regard to the patterns 
of human fingers will be the following: 

All fingers considered as a whole, we may look upon the ulnar loops as the 
typical human pattern, occurring on two thirds of all fingers and filling the 
place wherever this is not, for special reasons, occupied by other patterns. 
These reasons may be of two different kinds, conservative forces having 
preserved the phylogenetic oldest pattern, the whorl , in a certain percentage 
on all fingers but especially upon digits I and IV, and always more so upon 
the right hand than upon the left,—and on the other side adaptive forces 
influencing the reduction process of the pattern in correspondence with the 
function of each special finger, thus producing arches and radial loops upon 
the fingers in which the radial side is of special importance to their function, 
this is shown to be the case with digit V and III and, much more so, with 
the digit II. 


210 


KRISTINE BONNEVIE 


With regard to all pattern-types there is found a more or less conspicuous 
difference between right and left hands, the phylogenetically oldest patterns, 
the whorls being preserved in a higher percentage on right hands, especially 
on digits I, IV and V; and the younger, adaptive patterns, arches and radial 
loops being also more numerous on the right second finger than on the left. 

Phenomena of this kind may serve as a support of the assumption of 
Wilder (1916, p. 236) of an early adaptation of the right-handedness; it 
should, for an explanation the statistical facts mentioned, be supposed to 
have existed before the full inflation of the elevated pads upon digits I, 
IV and V, as well as before the present development of the adaptive patterns 
of digit II. 

Assuming that the different pattern-types which “are first seen in an 
embryo of approximately four months, since which age there is no indication 
of change throughout life ” (Wilder, 1916, p. 235), are determined through 
hereditary factors, a phylogenetic development from whorls into loops and 
arches may have been performed in two different ways, either through a 
loss of some factor determining the triradii of the whorls, or through the 
acquirement of new factors checking their development . A contrast should, 
in both alternatives, exist between individuals with their original, typical 
patterns practically intact, that is: with whorls on all fingers, and others 
in which the whorls are absolutely lost, and between these two extremes a 
series of transition stages. 

The heredity of the different stages of a gradual phylogenetic reduction of 
the whorl into less complicated patterns presents in itself, no special dif¬ 
ficulty to the understanding. A great difficulty arises, however, when one 
thinks of the adaptive results reached especially on the second finger through 
this reduction, an adaptation equally well fitting both a mechanical and a 
tactile function of the papillary ridges. As already mentioned by Wilder 
(1916, pp. 234-237) the adaptation of finger-patterns to their function is so 
much more peculiar, as the patterns themselves during individual life are 
not at all influenced by either age or function or other external forces. The 
pattern-types are determined long before birth, and cases like that of the 
two brothers with arches on all fingers (experiment 2, p. 209) strongly in¬ 
dicate that the determination is lying in the germ-cells. How, then, are 
these adaptive results reached? 

I can here only ask the question, and give no answer. But the answer 
once given, will be one of general importance applying also to phenomena of 
adaptation upon other and more important fields than that of finger- 
patterns. 


STATISTICAL INVESTIGATION OF FINGER PRINTS 


211 


REFERENCES 

F£r£, Ch. 1895 Note sur la sensibility de la pulpe des doigts. C. R. soc. biol., T. 47. 

Gallon, F. 1892 Fingerprints. London. 

Henry 1901 Classification and uses of finger prints. London. 

Johnson 1899 Pads on the palm and sole of the human foetus. Amer. Nat., vol. 33. 

Klaatsch 1887 tlber die Morphologie d. Tastballen. Anat. Ang., Bd. 2. 

Rlaatsch 1888 Sur Morphologie d. Tastballen d. Saugethiere. Morph. Jahrb., 
Bd. 14. 

Mohr, O. L. 1921 A case of hereditary brachyphalangy utilized as evidence in foren¬ 
sic medicine. Hereditas, Bd. 2. 

Retzius 1904 Sur- Kenntnis d. Entw. d. Korperform des Menschen wahrend d. foet- 
alen Lebensstufen. Biol. Untersuth. Neue Fig., Bd. 11. 

SCHLAGiNHAUi'EN 1905 Das Hautleistensystem der Primatenplanta. Morph. Jahrb., 
Bd. 33-34. 

Schlaginhauren 1906 Ueber das Leistenrelief d. Hohlliand und Fusssohlen-flache. 
d. Halbaffen, Affen u. Menschenrassen. Ergebn. Arch. u. Entw., Bd. 15. 

Whipple, Inez 1904 The ventral surface of the mammalian Chiridium, with especial 
reference to the condition found in man. Fubaehre. f. Morphol. u. Anthro- 
pol., Bd. VII. 

Wilder, H. H. 1916 Palm and sole studies I-VI. Biol. Bull., vol. 30. 

Wilder, H. H. 1919 Physical correspondence in two sets of duplicate twins. Journ. 
of Heredity, vol. 10. 


THE GENETICS OF FECUNDITY IN THE DOMESTIC HEN 
C. C. HURST 

Burbage , Leicestershire , England 

Five years’ experimental breeding on Mendelian lines from 1910 to 1916 
gave results which show that the fecundity of a hen depends on the com¬ 
bined action and reaction of many genetic factors. 

Seven main genetic factors have so far been identified in the experiments. 
Five of these, E, W, S, M, and H, influence fecundity, one affects egg-size 
(N) and one egg-color (C). Of the five factors influencing fecundity E and 
W together determine winter-production, S is manifested in spring- 
production, M influences autumn production, while H affects the produc¬ 
tion of any of the three periods. 

The seven pairs of Mendelian factors are as follows: 

E e Early and late sexual maturity of pullets. 

W w Fast and slow rate of winter production. 

S s Fast and slow rate of spring production. 

M m Slow and fast rate of autumn production. 

H h Broody and non-broody instinct (Hurst, 1905) 

N n Small and large egg-mode (Hurst, 1913) 

C c Brown and white egg-mode (Hurst, 1905-1913) 

The first of each pair is dominant and the second recessive. In order to 
analyze the complex and continuous data of egg-production a system of 
uniform gradings was formulated, which ultimately led to the identification 
of the above genetic factors. 

Sexual maturity was measured by age at first egg and graded in months of 
thirty days. 

Early grades (E) were 4 to 8 and late grades (e) 9 to 13, so that each hen 
had a somatic grading E4 to el3, while each hen and cock had a genetic 
formula EE, Ee, or ee. 

Rate of production was measured by percentage of eggs laid to number of 
days in lay, for each of the winter, spring, and autumn periods. These 
percentages were graded in tens. 

Winter period was from first egg to last day of February, spring period 
from March 1 to June 30 and autumn period from July 1 to last egg of year. 

212 


FECUNDITY IN THE DOMESTIC HEN 


213 


Fast winter grades (W) and fast spring grades (S) were 8 to 3 and slow 
winter grades (w) and slow spring grades (s) were 2 and 1. 

Slow autumn grades (M) were 0 to 2 and fast autumn grades (m) were 3 

to 6. 

Broodiness was measured by number of broody periods. Broody grades 
(H) were 6 to 1 and non-broody grade 0. 

Egg-size was measured by weight of egg in ten equal grades of J ounce 
ranging from lj to 2\ ounces. Small-egg grades were 1 to 4 and large-egg 
grades 5 to 9. The mode-grade of the first year’s eggs was taken to repre¬ 
sent the egg-size of each hen. 

Egg-color was measured by grading in 6 color shades of equal difference 
of intensity from dark-brown to chalky white, viz.: Dark brown (5), brown 
(4), light brown (3), cream (2), ivory (1), and white (0). Brown grades 5 
to 3 and white grades 2 to 0. The mode of the first year’s egg was taken to 
represent the egg-color of each hen. 

Four breeds were used in the experiments, namely, White Leghorn, 
White Wyandotte, Light Sussex and Pit Game, including three distinct 
utility strains of Leghorns and Wyandottes. 

In the course of the experiments more than 50,000 eggs were recorded, 
and each egg was weighed and graded for size and color. 

Four hundred and thirty pullets of four generations were reared from 
mated pairs, each pullet was tested for egg-production for one year after 
first egg where possible. 

The unequal influences of the external conditions were minimised by the 
adoption of certain uniform methods of housing, feeding and exercise which 
involved the keeping of each individual in a single house and run. The 
factorial constitutions of 42 sires and 135 dams for the seven genetic factors 
were determined as far as possible. 

The genetic constitutions of the birds were ascertained by a combination 
of the following four tests: (1) Somatic (hens only), (2) parental, (3) 
progeny qualitative, (4) progeny quantitative. This combination method 
of analysis was necessary owing to the comparatively small families of 
pullets raised from the mated pairs and proved to be critical in its complex 
harmonies. 

The following table gives a summary of the results of the experiments 
with White Leghorns and White Wyandottes on the basis of the scheme of 
the seven pairs of factors and the genetic analysis involved. 

Reduced to terms of dominants and recessives the total numbers are 
1195: 687R showing a net deficiency of 17.75 dominants and a correspond¬ 
ing excess of recessives in the total of 1882 observations, or less than 1 per 
cent. 


214 


C. C. HURST 


The gross divergence in the results of the seven factors amount to 29.25 
or less than 2 per cent. 


Summary of results in White Wyandottes and White Leghorns 


CHARACTERS 

TOTALS 

DOMINANT 

OB¬ 

SERVED 

EX¬ 

PECTED 

RECESSIVE 

OB¬ 

SERVED 

EX¬ 

PECTED 

Sexual maturity .. 

335 

Early (E) 

286 

289.00 

Late (e) 

49 

46.00 

Winter rate. 

266 

Fast (W) 

231 

231.75 

Slow (w) 

35 

34.25 

Spring rate. 

224 

Fast (S) 

216 

217.25 

Slow (s) 

8 

6.75 

Autumn rate. 

194 

Slow (M) 

140 

145.50 

Fast (m) 

54 

48.50 

Broodiness. 

201 

Broody (H) 

50 

58.50 

Non. B. (h) 

151 

142.50 

Egg-size. 

331 

Small (N) 

135 

128.75 

Large (n) 

196 

202.25 

Egg-color. 

331 

Brown (C) 

137 

142.00 

White (c) 

194 

189.00 

Totals. 

1882 


1195 

1212.75 


687 

669.25 


Eighteen definite exceptions appeared of which 2 proved to be somatic 
and not genetic, 10 were slight exceptions probably of the same nature 
3 were pathological, 2 were possibly incomplete dominants, while 1 was 
apparently a true mutation. 

Pearl’s discovery of two genetic factors for winter production in Plym¬ 
outh Rocks (1912) confirmed by Goodale and MacMullen in Rhode 
Island Reds (1918 and 1919) is also confirmed in White Leghorns and 
White Wyandottes. But as neither Pearl’s factors Li and L 2 nor Goodale’s 
factors A and B are individually identified with sexual maturity (E) or 
with rate of production (W), and as Pearl’s factor L 2 is sex linked, while the 
others are not, it seems impossible to identify factors E and W definitely 
either with Pearl’s Li and L 2 , or with Goodale’s A and B. No sex linkage 
was found either in the White Leghorns or White Wyandottes used in the 
experiments. In this respect these two breeds agree with Goodale’s Rhode 
Island Reds rather than with Pearl’s Plymouth Rocks. 

No winter zero birds appeared in the White Leghorns, though 9 appeared 
in the White Wyandottes, and the White Leghorns. All were late maturers 
(ee) and all the five zeros tested genetically carried the factor W, so that 
Pearl’s interpretation that the zero winter producers in Plymouth Rocks are 
mostly due to the absence of both production factors is not borne out in 
Leghorns or Wyandottes. The Wyandotte zeros on the contrary resemble 
Goodale’s zeros in Rhode Island Reds, their zero production being due 
entirely to their very late sexual maturity. It is possible, however, that this 
very late maturity may be due to a genetic sub-factor. 

In all the factors except H (broodiness) dominance tends to be incomplete, 
and the heterozygotes are more or less intermediate. In many cases it is 




























FECUNDITY IN THE DOMESTIC HEN 


215 


possible to recognize the homozygous individuals by their extreme grades, 
though not always. 

In each of the seven main genetic factors for egg-production there are 
distinct indications of the presence of one or more sub-factors, but the 
numbers are insufficient to determine these and their allelomorphs satis¬ 
factorily. It is possible that an intensive study of a single set of these 
sub-factors would throw some light on the nature of the “fractional factors ” 
of Bateson (1914) and the so-called “multiple allelomorphs” of Sturtevant 
(1913) and others. 

There appears to be a definite difference of rhythm between the fast 
(W) and fast (S) birds and particularly between the discontinuous slow 
(w) and the discontinuous slow (s) birds, and it is not improbable that the 
slow (s) birds are pathological. 

The M factor is clearly of a different nature from the factors W and S, 
seeing that in the two latter fast rate is dominant, while in the former it is 
recessive. A striking somatic difference was observed between M and m 
birds apparently coinciding with their different rates of production. The 
M birds are deep autumn moulters and slow producers, while the m birds 
are partial autumn moulters and fa^t producers. The m mutation, which 
has incidently made the “300-egg hen” possible, is interesting genetically 
on account of its comparatively recent appearance. 

Owing to the fact that a sensible proportion of broody hens do not show 
their broodiness until their second season of laying, it has not been possible 
to ascertain the true nature of the “non-broodies” in the experiments, as 
most of them were only observed during their first laying year. Conse¬ 
quently the results show a deficiency of broodies. The appearance of a few 
broody exceptions in the non-broody Leghorns gives support to Punnett’s 
(1920) suggestion of the possible presence of an inhibitor to the broody 
factor in certain non-broody birds. The evidence for the HI scheme for 
broodiness is admittedly incomplete, and experiments on a considerable 
scale would be necessary to demonstrate it. At the same time it seems to 
bring into line most of the complicated and conflicting data of the genetics 
of broodiness that have been published. 

Evolutionary significance. The genetic results indicate the gradual 
evolution of the increase of fecundity in the hen by a succession of definite 
and discontinuous steps or mutations. Three of these mutations are 
dominant, viz.: early maturity (E), fast winter rate (W) and fast 
spring rate (S), while two are recessive, viz: fast autumn rate (m) and 
probably non-broodiness (h). A single case of the solitary appearance of 
a recessive mutation was observed, in the experiments, which apparently 


216 


C. C. HURST 


originated in a male Wyandotte by the loss or absence of the factor N 
(small-egg mode) in a single gametogenesis giving rise to a daughter and 
granddaughters with large-egg modes (n). If the original mutation (n) 
arose in the same way in the wild species, at least two generations would 
elapse before it could appear owing to its recessive nature, and the lack 
of a recessive mate. In view of the recent contributions to the chromosome 
theory of the mechanism of heredity by Morgan (1919) and his colleagues, 
it is unfortunate that the above experiments do not provide many suitable 
matings for testing linkages satisfactorily, and investigations in this direc¬ 
tion might lead to results of considerable genetic and economic importance. 

Economic significance. It is obvious that these genetic results are of 
considerable economic importance, and the possibilities of their application 
to practical poultry breeding will be dealt with elsewhere. 

Eugenic significance. From the problem of the fecundity in the domestic 
hen to the problem of fertility in man is admittedly a far cry. The cases 
are by no means parallel though they are in many respects analogous. 

Pearl and Surface (1909) rightly distinguish fecundity from fertility, 
and use the term fecundity to ‘‘designate the innate potential reproductive 
capacity of the individual organism , as denoted by its ability to form and 
separate from the body mature germ-cells. Fecundity in the female will 
depend upon the production of ova, and in the male upon the production 
of spermatozoa. In mammals it will obviously be very difficult, if not im¬ 
possible, to get reliable quantitative data regarding pure fecundity. On 
the other hand we would suggest that the term “fertility” be used to desig¬ 
nate the total actual reproductive capacity of pairs of organisms, male and 
female, as expressed by their ability when mated together to produce 
(i.e., to bring to birth) individual offspring. Fertility, according to this 
view, depends upon and includes fecundity, but also a great number of other 
factors in addition. Clearly it is fertility rather than fecundity which is 
measured in statistics of birth of mammals.” 

We have seen above that at least five pairs of genetic factors are concerned 
with fecundity in the domestic hen, and if we accept Pearl’s view that 
fecundity is only a part of fertility, we may expect fertility in fowls to be of 
a still more complex nature genetically. But the genetic factors of fertility, 
are only a part of the general problem of fertility, for to these must be added 
the multitudinous somatic or environmental factors, many of which are yet 
unknown to science. 

If the problem of the causes of fertility in poultry is clearly so complex, 
one can hardly expect the problem of the causes of fertility in man to be of 
a more simple nature. 


FECUNDITY IN THE DOMESTIC HEN 


217 


It is evident that there can be no eugenic control of fertility in man 

until the causes of that fertility are known, and the first line of attack 

might well be directed to the identification of the genetic factors of human 

fertility and sterility. 

REFERENCES 

Bateson, W. 1914 Presidential address to British Association for the Advancement of 
Science (Australia). 

Goodale, H. D. 1918 Internal factors influencing egg-production in the Rhode Island 
Red breed of domestic fowl. Ameri. Nat., lii, 65, 209, and 301. 

Goodale, H. D., and MacMullen, Grace 1919 The bearing of ratios on theories of 
the inheritance of winter egg-production. Jour. Exper. Zool., xxviii, April, 
pp. 83, 124. 

Hurst, C. C. 1905 Experiments with poultry. Reports to the Evolution Committee of 
the Royal Society, Report 11, pp. 131-154. 

Hurst, C. C. 1913 Breeding experiments with utility poultry. Report V to Birming¬ 
ham Meeting of the British Association for the Advancement of Science. 
(Sections D, K and M.) 

Morgan, T. H. 1919 The physical basis of heredity. (Lippincott.) 

Pearl, R. 1912 The mode of inheritance of fecundity in the domestic fowl. Jour. 
Exper. Zool., xiii, August, pp. 153-268. 

Pearl, R., and Surface, F. M. 1909 Data on the inheritance of fecundity obtained 
from the records of egg-production in the daughters of “200-egg” hens. 
Maine Agricultural Experiment Station Annual Report, Bulletin 161, pp. 
49-84. 

Punnett, R. C., and Bailey, P. G. 1920 Genetic studies in poultry. II. Inheritance 
of egg-color and broodiness. Jour. Genetics, December, x, pp. 277-292. 

Sturtevant, A. H. 1913 The Himalayan rabbit case, with some considerations on 
multiple allelomorphs. Amer. Nat., xlvii. 


INHERITANCE OF MENTAL DISEASE 

A. MEYERSON 
Boston , Massachusetts 

I. INSANITY IS NOT A UNIT AND AS A TERM HAS NO PLACE IN THE SCIENCE 
OE MENTAL DISEASES 

Like many another word insanity exercises a baleful influence over our 
thought. Originating in the idea that all mental disease was of the same 
general nature, and having its roots partly in the superstition that the 
insane persons are possessed, and partly in the legal concept of insanity as a 
condition in relation to the ability to care for one’s property and one’s 
social obligations, it has caused a marked confusion in psychiatric thought 
and especially in relation to the transmission from generation to generation. 

Mental diseases are as different from one another as diseases of the re¬ 
spiratory tract. We recognize that tuberculosis is biologically an entirely 
different entity from the pneumonia caused by the pneumococcus, and that 
both differ from asthma, and that syphilis of the lung is in no wise related 
biologically except as to place of incidence with the previously mentioned 
diseases. Similarly with mental diseases there are types that are related 
to injury or disease of the brain, and the injured or diseased conditions vary 
biologically to a marked degree. Thus one of the commonest diseases is 
general paresis, which is caused by the Spirochaeta pallida or the organism 
of syphilis. This has no conceivable biological relationship to the mental 
disease caused by tumor growth. In fact, a tumor of the brain is biologi¬ 
cally more related to cancer of the breast than it is to the rest of the subject 
matter of psychiatry. Trauma or injury to the brain causes mental disease 
but it is absurd to link this up with general paresis or tumor. Further, one 
finds mental disease in relation to meningitis and tuberculosis, and these con¬ 
ditions are related not so much to the other mental diseases but to the 
general problems of infection and its spread, and tuberculosis. In fact, 
psychiatry is a sort of nodal point into which a great many roads lead. 

There are mental diseases caused by direct toxic influences. For example, 
alcohol caused somewhere from 15 to 20 per cent of all cases of “insanity. ” 
Morphine has its own type of psychosis, as have cocaine and heroin. 
Further, there are different mental diseases associated with toxemias of 

218 


INHERITANCE OF MENTAL DISEASE 


219 


pregnancy and with toxemias occurring in the course of other infectious 
diseases such as influenza, pneumonia, typhoid, etc. Then there are 
mental states of an abnormal kind and which become in many cases 
committable to insane hospitals, which directly follow great exhaustion. 
All of the above named groups of diseases are classed as toxic exhaustive 
psychosis. The heredity of these groups is nil or almost nil, as the fig¬ 
ures from the Taunton State Hospital show. 

There is a group of mental diseases of unknown origin, probably not re¬ 
lated to any definite pathology of the brain at all, more likely related to 
disease of the endocrinal glands, and concerning which there is a remarkable 
diversity of opinion as to cause. For example, one group of observers think 
they have their origin in psychological traumata and trace them back to 
complexes of one type or another; others believe they arise in personality 
trends which become accentuated as time goes on into true mental disease; 
still others link them up with a disturbance of the endocrines, and another 
group tries to bring back their pathology to actual disease of the brain. 
In fact, no one knows their pathology, and this group is the only group of 
importance insofar as heredity of mental disease is concerned. This group 
comprises dementia praecox, manic depressive insanity and the involution 
psychosis, including certain of the diseases found in old age. As a matter 
of fact, even these diseases are not, biologically speaking, entities. There is 
at the present time great dissatisfaction in psychiatry with the scheme of 
classification by which they are designated as above. Take for example 
dementia praecox, one of the most common of mental diseases and isolated 
as an entity by the German psychiatrist Kraepelin. Ten years ago it was 
hailed as a triumph; today there are few who believe that this concept is 
anything more than a grab-bag into which many diseases are stowed. 

II. CERTAIN OF THE STATISTICAL PHENOMENA RELATING TO MENTAL DISEASE 
The neuropathic heredity of the insane and non-insane 

This question has not been personally studied. The conclusions here 
stated are taken from the only two important studies on the subject, those 
of Koller and Diem. A few of their data are to my mind clear-cut enough 
to be important. 

1. There is very much general neuropathic heredity in the direct and 
collateral relatives of both the insane and sane. 

2. There is far more insanity in the families of the insane, and this is 
especially true of the parents and grandparents of the insane. 


220 


A. MEYERSON 


3. Insane uncles and aunts occur about as frequently in the families of 
the sane as in those of the insane; therefore, collateral insanity is of relatively 
little importance unless associated with parental insanity. 

4. The sane seem to have as much, or even more, of nervous disease, senile 
dementia, and apoplexy in their ancestry. This would throw out of court 
as useless the questions as to nervous disease, apoplexy, etc., in insane 
hospital histories. It would entirely vitiate the value of such works as 
Davenport, in which apoplexy is considered a neuropathic taint when, as a 
matter of fact, the question is entirely arterio-cardio-renal. 

The work of Koller and Diem is vitiated to a certain extent by the fact 
that both have taken insanity as a sort of unit and have regarded normality 
as something fixed. Nevertheless, their facts are of importance and 
similar studies need be done more scientifically and on a larger scale. 

The marriage rate of the insane 

In the four groups studied personally, including males and females— 
alcoholic insanities, general paresis, dementia praecox, and senile dementia— 
we find, first, that the males in the alcoholic paretic and dementia-praecox 
groups marry less than do the females. In the seniles, although the per¬ 
centage of married men is greater, the totals of those who have entered 
conjugal relations at one time or another are about equal. Looking some¬ 
what closer, it is found that in paresis there is only a slight difference in 
favor of the female, while in dementia praecox this difference is much 
further increased; thus, if these groups may be held to constitute a menace 
by virtue of their ability to transmit the psychotic taint to another genera¬ 
tion, the female of the species, to use a well known phrase, is more dangerous 
than the male. 

The seniles and the general paretics marry but slightly less than do the 
same age groups in the total population; the alcoholics show a decided 
falling off as compared with the total population; while the male dementia 
praecox has an exceedingly low marriage rate. That is, whatever is back 
of dementia praceox, it operates against self-perpetuation. Something of 
the same internal mechanism is seen in the case of alcoholism. This 
mechanism operates very little, if at all, in the case of paresis and the 
senile psychoses. One might conclude that if there is an inborn defect in 
these diseases it is by far greatest in dementia praecox, is next in alcoholic 
insanity, and least of all in syphilis and the senile psychoses. 

We are shown that marriage acts as a barrier to the propagation of the 
abnormal insofar as this is connected with endogenous factors. It is not 
a barrier against certain of the exogenous race poisons, such as, for example, 


INHERITANCE OF MENTAL DISEASE 


221 


syphilis, at least in that form which leads to paresis. We need to strengthen 
the barrier against the endogenous diseases, as for example, dementia 
praecox, but not nearly so much as we need to strengthen it against the 
exogenous, as, for example, syphilis. 

General statistics 

For the purpose of comparison, figures obtained by Mott in his studies 
have been included in the following summary. 

1. More females than males are concerned in family psychoses. The 
studies at the Taunton State Hospital show 808 females and 739 males. 

2. The mother-daughter group is much more common than the father- 
daughter group—80 to 59, as shown by the Taunton figures; 137 to 103, 
as shown by Mott’s figures. 

3. The mother-son groups are about equal. 

4. Mother-son and father-son are about even—56 to 59, in accordance 
with Taunton figures; 96 to 88, as shown by Mott’s figures. 

5. Sisters alone decidedly outnumber brothers alone—80 to 57 according 
to Taunton figures; 211 to 140, according to Mott’s figures. 

If insane women transmit their mental peculiarities to their female 
children more than they do to their male, then the greater marriage rate 
amongst insane women may decidedly play a part in determining the 
preponderance of insane women. Furthermore, men migrate more than 
women, and so in any given hospital district the female descendants of 
insane ancestors would be more apt to appear in the asylum than the male 
descendants even if given equal rate of incidence. That is to say, a larger 
part of these men would end in jails or in hospitals in districts remote from 
their former homes, etc. 

Anticipation or antedating 

This term is used (Darwin) to describe the earlier appearance of mental 
disease in the younger as compared with the older generation. This phenom¬ 
enon has been given great prominence by Mott and his co-workers. He 
regards it as an effort of nature to get rid of the disease by crystallizing it 
in a few descendants and making them more easily vulnerable or unfitted 
to propagate by being brought early to hospitals. There are thus in Mott’s 
definition of anticipation, first, a crystallization of the insane elements, 
leaving other descendants free from disease, and second, earlier onset of the 
psychosis in the affected members. 

All of the figures, and especially in the second-generation cases, seem to 
point clearly to the validity of the phenomenon of antedating or anticipa- 


222 


A. MEYERSON 


tion. A closer examination, however, shows one great fallacy underlying 
the statistics and that is as usual, the method of collection. In a majority 
of cases the period of time during which the cases have collected is not clear 
(for example, in Mott’s statistics, where no mention is made of the period 
during which his figures have been collected), but it is generally within 
thirty years. As the average ancestor is at least that much older than his 
average descendant, it would be practically impossible for the descendant 
to be older than the ancestor at time of commitment, and the age difference 
between ancestory and descendant would generally be from thirty-three 
years to zero, in favor of the ancestory, and in certain cases where the de¬ 
scendant enters the hospital before the ancestor it would be thirty years or 
more. This means that in the cases of some of the investigators it has been 
possible for the younger descendants to enter the hospital, but time enough 
has not elapsed for the older descendant to arrive. As the Taunton figures 
have been collected since 1854, this objection does not obtain with such 
force. Nevertheless, the rate of commitment has increased within the last 
generation so that even in my cases the bulk of families has arrived within 
the last thirty years and thus the factor stated above still plays its part in 
falsifying the manifest results. 

It is probable, therefore, when all is said and done, that in a very large 
proportion of cases the descendants of the insane who themselves become 
insane do so at an earlier age than their ancestors. Moreover, as is well 
known, these descendants have a much lessened chance for marriage and 
so this factor of anticipation seems to be, as Mott believes, a potent factor 
for race regeneration through elimination. One need not, of course, sub¬ 
scribe to any such anthropomorphic absurdity as that nature has any inten¬ 
tions in the matter. It is perhaps better to speak of this phenomenon as the 
downward trend of psychiatric families, just as the same descendants repre¬ 
sent their upward trend. 

The transmission of mental diseases is practically nil insofar as the organic 
disease of the brain goes. It may be stated on the basis of personal research 
that there are very few families in which mental disease occurs generation 
after generation in anything like the way other hereditary characters are 
transmitted. At the Taunton Hospital there are records of one four- 
generation family; there were eighteen families in which three generations 
were represented on the records. In the majority of these cases the relation¬ 
ship of the mental disease of the grandparent to the mental disease of the 
grandchild was in great doubt, and in fact, in the majority of cases there was 
a much clearer relationship to some factor entering with a new line of 
ancestors. In other words, it is rare that one can see a direct relationship 


INHERITANCE OF MENTAL DISEASE 


223 


between a mental disease in a grandparent and one in a grandchild. The 
mental diseases Which occur in two generations are very common. On 
the whole the following may be stated. Other mental diseases besides 
dementia praecox, manic depressive insanity, and the involution psychoses 
rarely, if ever, can be shown to have a direct hereditary value. This is 
true of the alcoholic psychoses. That is to say, the fact that an ancestor 
has a mental disease due to alcoholism does not seem to predispose his 
descendant to mental disease of other type. A fairly uniform relationship 
can be shown between the mental disease of the ancestor, the mental 
disease in the parent, and the mental disease in the child somewhat along 
these lines (if the child is subject to mental disease for it needs here be 
emphasized that the great majority of the descendants of this class of the 
insane do not themselves become insane): 

1. Given a paranoid type of psychosis, including in this term true para¬ 
noia, paranoid dementia praecox, and unclassified paranoid states, what 
types of mental disease may we expect in the descendants? The answer is 
definitely that we may expect either paranoid disease or ordinary dementia 
praecox in the insane descendants, and this statement is corroborated by 
Jolly, Luther, Krueger, Rosanoff, and Albrecht. Practically no manic- 
depressive descendants appear. 

2. Given well defined dementia praecox in an ancestor, what type of 
mental disease may we expect in the descendant? Dementia praecox 
follows in the large majority of cases. Feeblemindedness of a type which 
is perhaps very early dementia praecox is relatively common. Epilepsy 
is noted. Manic-depressive insanity occurs very rarely if at all. 

3. Given a well defined manic-depressive insanity in an ancestor, what 
type of mental disease may we expect in the descendants? There appear 
to be two main trends, one toward manic-depressive insanity and the other 
toward dementia praecox. Difficulties in diagnosis play a large part in 
solving this particular question, but it is certain that short attacks of 
mental disease bearing the earmarks of manic-depressive in an ancestor 
are followed in insane descendants by a definite dementing psychosis 
corresponding to what this uncertain generation calls dementia praecox. 

4. Given an ancestor with involution psychosis, what type of mental 
disease may we expect in the insane descendant? It seems to be universal 
experience that dementia praecox follows involution psychosis and that 
apparently no matter what the type of the involutional insanity. 

5. Given an ancestor with senile psychosis, what type of mental disease 
may we expect in the insane descendant? The Taunton figures, as well as 
those of Vorster, Albrecht, Jolly, Luther and Krueger, all find that what is 


224 


A. MEYERSON 


termed “senile dementia” is of such heterogeneous nature that what might 
be expected, occurs; that is, dementia praecox, manic-depressive, paranoid 
psychoses, imbecility and epilepsy, all are found in the insane descendants. 

If, then, we survey the facts which have here been presented together 
with the trends observed in the literature, we find that the paranoid and 
catatonic diseases trend finally to dementia praecox; that manic is succeeded 
by manic and in a varying proportion of cases by dementia praecox; that the 
senile and involutional psychoses, if paranoid, or more properly involution 
and senile, trend towards paranoid diseases and dementia praecox. Manic 
states of the senium follow the rule of manic states elsewhere. Neither for 
organic brain disease nor alcoholic psychoses can anything very definite be 
said, except that in the cases studied, wherever adequate history has been 
obtained, the psychosis in the descendant of the alcoholic or of the patient 
with organic brain disease can be related to some other more definite psycho¬ 
pathic feature than either alcoholism or organic brain disease. It will thus 
be seen that all roads seem to lead to dementia praecox and from thence to 
imbecility. 

With regard to the horizontal transmission of insanity, it may be thus 
restated: Given an insane person, what psychosis may we expect in his 
insane brothers or sisters? 

1. The psychoses of brothers and sisters trend on the whole to be of 
similar type. 

2. If one brother or sister has dementia praecox, the chances for de¬ 
mentia praecox in any other insane brother or sister are very great. Feeble¬ 
mindedness, or what is called feeblemindedness, occurs frequently. This, 
as has been before stated, may merely be congenital dementia praecox with 
dementia as the leading feature. 

3. Manic-depressive insanity and dementia praecox are said to occur to¬ 
gether in the same family group of brothers and sisters. Such cases are 
usually atypical, and really clear-cut cases with such association are rare. 

III. GENERAL REMARKS 

This paper has attempted to emphasize in a very brief manner the fact 
that mental diseases are not unitary and that each type of mental disease 
must be studied by itself in order to discover its hereditary relationships. 
Most of the mental diseases have no hereditary relationships; a few as stated 
above do. Any individual whose candidacy for matrimony is being con¬ 
sidered need not hesitate to marry if his ancestors have died in insane hos¬ 
pitals of the organic brain diseases, of alcoholic insanity, and of those groups 
which are non-hereditary, but the situation is entirely different when those 
diseases which have been discussed above are concerned. 


INHERITANCE OF MENTAL DISEASE 


225 


We are very far from establishing the fact that we are dealing with biologi¬ 
cal entities concerning which we may make any kind of general conclusions 
in relation to heredity. The insane have normal descendants; normal folk 
have insane descendants in a perfectly bewildering and inexplicable fashion. 
When all the facts are gathered in an impartial manner this is the one phe¬ 
nomenon that stands out. It is more likely in my opinion that we are deal¬ 
ing with a disease of the germ-plasm and injury inflicted upon it, which is 
thereupon transmitted for a long or short time through one or more genera¬ 
tions according to the severity of the injury. There is plenty of evidence at 
present collected to show that germ-plasms can be so injured and that the 
disease may thereupon appear without further injury, be inflicted on succes¬ 
sive generations, and in fact grow worse as time goes on. The work of 
Stockard and his associates, the work done in transmitting diseases of the 
lens, and certain evidences adduced by Forel and his associates concerning 
what they call blastophoric influence is of high significance. As eugenists 
we must not blind ourselves to the fact that stocks may become sick in the 
same sense that individuals do; that this sickness may pass down for several 
generations and then the stock recover exactly as does an individual from 
an infection, though of course the stock may die just as the individual 
does. The most fertile work that can be done at present in the study of 
the inheritance of mental disease is to look for the agents that injure germ- 
plasms rather than try to link up the transmission of mental disease with 
the phenomena of Mendelism or other great biological laws. In other words 
the problem of the transmission of mental disease is a clinical medical prob¬ 
lem to be studied by laboratory methods as in the rest of experimental 
medicine. 


INHERTIANCE OF MENTAL DISORDERS 
AARON J. ROSANOFF 

Kings Park State Hospital , Kings Park , Long Island , New York 


The object of this communication is to present a summary of the more im¬ 
portant known facts concerning the inheritance of mental disorders and to 
attempt a synthetic treatment of them. 

Mental disorders consist of a series of more or less sharply defined clini¬ 
cal entities on the causation of which variable relative parts are played 
respectively, by inborn and environmental factors. Numerically the most 
important group consists of those disorders which have, by a sort of common 
consent, been designated constitutional , the implication being that the inborn 
factors play an essential part in their causation. It is with this group that 
my paper deals. 

The principal clinical entities included are: mental deficiency, epilepsy, 
dementia praecox, paranoia, manic-depressive psychoses, psychoneuroses, 
and psychopathic personalities. A more detailed classification of them is 
given in the following chart. 


Mental deficiency. 


Idiocy 

Imbecility 

Moronism 

Borderline intelligence 


Epilepsy.... 


{ Epileptic personality 
Psychotic episodes 
Epileptic deterioration 


Paranoia 


Various monomanias 


Dementia praecox. 


Manic-depressive psychoses. 


Simple form 
Hebephrenic form 
Catatonic form 
Paranoid form 

Depressed form 
Manic form 

Mixed and alternating forms 
Involutional forms 


226 









INHERITANCE OF MENTAL DISORDERS 


227 


Psychoneuroses. 


Psychopathic personalities. 


j Hysteria 
■j Neurasthenia 
[Psychasthenia 

Inadequate personality 
Paranoid personality 
Emotional instability 
<{ Criminalism 
Pathological lying 
Sexual psychopathies 
Nomadism 


Institutional experience shows that the constitutional mental disorders 
run in families, and special statistical studies point to heredity as being the 
origin of psychopathic constitutions (1, 2, 3). The latter fact is today no 
longer in question; but there is considerable difference of opinion as to the 
relative amounts of causation attributable to heredity and to other factors. 

The obvious reason for this difference of opinion lies in our inability to 
measure exactly the factors involved. Rough measurements are, however, 
possible, and accumulated evidence seems to be to the effect that the rela¬ 
tive importance of the factor of heredity varies in individual cases, first, 
with the clinical subdivision, and, second, within each subdivision, with 
age of onset, amount of demonstrable external cause, intensity of manifes¬ 
tations, their intractableness or persistence in spite of treatment, etc. 

Thus, mental deficiency and, to a lesser extent, epilepsy are subdivisions 
in which heredity alone in most cases determines the manifestations. On 
the other hand, psychoneuroses, especially hysteria, most often develop in 
reaction to special external situations, being, under usual conditions, latent. 
Other subdivisions hold, in this respect, intermediate positions. 

Within the same subdivision, say that of dementia praecox, are found, on 
the one hand, cases which present abnormal psychic traits all through child¬ 
hood, onset of grave psychotic symptoms at puberty or before in the absence 
of pathogenic environmental influences, and rapid passage into chronicity 
and deterioration in spite of all efforts of treatment; and on the other hand, 
cases in men of apparently normal psychic make-up, who, perhaps at middle 
age, following financial and marital troubles, take to heavy drinking and 
develop an acute hallucinosis or some other schizophrenic episode, which 
subsides under no other treatment than a few days’ rest in a hospital. 

A highly significant fact is that of frequently observed atavistic heredity: 
after one or more generations in direct line of descent have been skipped, an 
ancestral mental disorder reappears, sometimes affecting two or more in¬ 
dividuals in a sibship. From a Mendelian standpoint this is, of course, 






228 


AARON J. ROSANOFF 


suggestive of recessiveness in relation to normal mental constitution. This 
seems to be true of all constitutional mental disorders, regardless of variety, 
with the possible exception of the manic form of manic-depressive psychoses 
(4,5,6,7). 

Another significant fact is that of dissimilar heredity. All psychopathic 
members of a given family do not necessarily suffer from the same clinical 
form of mental disorder. More often than not they differ from one another 
either qualitatively, i.e., in respect of clinical variety, or quantitatively, i.e., 
in respect of severity of the disorder (8, 9, 10, 11, 12). 

The manner of distribution of the different clinical varieties of mental 
disorders in a family is apparently not a random one. Many studies have 
resulted in the observation that, although manic-depressive ancestors not 
infrequently have dementia praecox descendants, the reverse is very rare 
(13, 14, 15, 16). 

Based upon this and other similar observations a theory has been sug¬ 
gested, according to which the various psychopathic conditions possess 
different degrees of recessiveness and may be ranged in a scale of dominance 
in the following order: normal condition, manic-depressive psychoses, 
dementia praecox, epilepsy (7, 11). 

The fact of some sort of relationship of the clinically distinguished enti¬ 
ties to one another is suggested not only by family studies, but also by the 
existence of transitional and borderline cases presenting mixed manifesta¬ 
tions: dementia praecox with epileptiform convulsions; manic-depressive 
psychoses with an admixture of catatonic, delusional, hallucinatory, or other 
schizophrenic symptoms; psychoneuroses with occasional elated or de¬ 
pressed spells, etc. (11, 17, 18, 19, 20). 

The exact definition and measurement of the several clinical entities are 
rendered difficult by the fact of the great variations in their manifestations 
in different cases and in the same case at different times. These variations 
are spoken of among medical men as equivalents. Progress in psychiatry 
in the past two or three decades has been marked by a simplification of 
classification through a far reaching extension of the conception of clinical 
equivalents; and it would seem that a further extension of this conception is 
indicated by new accumulations of material (7). 

What bearing has our accumulated knowledge of the part played by 
heredity in the production of mental disorders on applied eugenics? Would 
there be justification in advocating restriction of propagation in all cases in 
which signs of constitutional mental disorder can be definitely established? 

In this connection it is sometimes forgotten that the aim of eugenics is 
not only to prevent the propagation of socially undesirable traits, but also 


INHERITANCE OF MENTAL DISORDERS 


229 


to conserve and stimulate the propagation of socially desirable ones. The 
great majority of individuals, insane as well as sane, present combinations 
of desirable and undesirable traits in endless variety. A general judgment 
seems hardly possible; each case must receive special consideration. The 
presence of a mental disorder in gravest degree is not incompatible with 
intellectual activity of the highest order and value to society, as is witnessed 
by the cases of Jean Jacques Rousseau, Gustave Flaubert, William Cowper, 
Auguste Comte, Julius Robert Mayer, Foodor Dostoyovski, and many 
others (21). 

Moreover, judgment as to desirability or undesirability of a given trait 
is apt to vary in different countries, or parts of countries, or, in the same 
country, at different times, depending on type of civilization and prevailing 
social ideals and standards. It would seem to be a part of a healthy con¬ 
servation to refrain from the employment of any eugenic measure which is 
irrevocable—such as sterilization—at least for the present. Against segre¬ 
gation, on the other hand, there is no such objection. It meets, indeed, 
not only the requirements of eugenics, but also other social requirements, 
and should be even more freely practiced than it is. 

Segregation is a measure which requires no forcing. Existing institutions 
are greatly overcrowded and many of them have long waiting lists. It 
has been estimated that even in a state like New York, which stands among 
the highest in respect of providing facilities for segregation, such facilities 
would have to be doubled before the existing demands could be met (22). 

On the scientific side the most urgent need seems to be two-fold, (1) to 
define more clearly the various qualitatively distinguishable psychopathic 
traits, including a fuller knowledge of clinical equivalents, (2) to devise 
for temperamental traits methods of more precise measurement, as has 
already been done for intellectual traits. 

REFERENCES 

(1) Roller: Beitrag. zur Erbfichkeitsstatistik der Geisteskrankheiten im Kanton 

Zurich. Vergleichung derselben mit der iiblichen Belastung gesunder Men- 

schen durch Geistesstdrungen und dergleichen. Arch. f. Psychiatrie, 1895, 

27, p. 268. 

(2) Diem, O.: Die psycho-neurotische erbliche Belastung der Geistesgesunden und der 

Geisteskranken. Arch. f. Rassen- u. Ges., Biol., 1905, ii. 

(3) Heron, D.: A first study of the statistics of insanity and the inheritance of the 

insane diathesis. Eugenics Laboratory Memoirs, II. 1907. 

(4) Goddard, H. H.: Heredity of feeble-mindedness. Bulletin No. 1, Eugenics 

Record Office, 1911. 

(5) Goddard, H. H.: The Kallikak Family. A study in the heredity of feeble¬ 

mindedness. New York, 1912. 


230 


AARON J. ROSANOFF 


(6) Davenport and Weeks: A first study of inheritance , of epilepsy. Bulletin 

No. 4, Eugenics Record Office, 1911. 

(7) Rosanoff and Orr: A study of heredity in insanity in the light of the Mendelian 

Theory. Bulletin No. 5, Eugenics Record Office, 1911. 

(8) Vorster: Ueber die Vererbung endogener Psychosen in Beziehung zur Klassikation. 

Monats. f. Psych, u. Neurol., 1901, ix. 

(9) Berse, J.: Die hereditaren Beziehungen der Dementia praecox. Leipsic and 

Vienna, 1910. 

(10) Berze, J.: Die manisch-depressive Familie H. Monats. f. Psych, u. Neurol., 

1909, xxvi. 

(11) Rosanoff, A. J.: Dissimilar heredity in mental disease. Amer. Journ. of Insanity, 

1915, lxx. 

(12) Moore, A. S.: Some preliminary observations concerning the types of psychoses 

occurring in the individual members of families. N. Y. State Hosp. Bul¬ 
letin, May, 1913. 

(13) Myerson, A.: Psychiatric family studies. Amer. Journ. of Insanity, 1917 and 

1918, lxxiii and lxxiv. 

(14) Jolly, P.: Die Hereditat der Psychosen. Arch. f. Psych, u. Nervenkrank., 1915, lii. 

(15) Krause: Ueber Vererbung von Gesiteskrankheiten. Allg. Zeitschr. f. Psychiat., 

1903, lx. 

(16) Rudin, E.: Zur Vererbung undV Neuentstehung der Dementia praecox. Mono- 

graphien aus dem Gosamtgebiete der Neurol, u. Psychiat., 1916, 12. 

(17) Kraepelin, E.: Dementia praecox and paraphrenia. (English translation by 

R. M. Barclay.) Edinburgh, 1919. 

(18) Bleuler, E.: Dementia praecox oder Gruppe der Schizophrenien. Leipsic and 

Vienna, 1911. 

(19) Simon, T. W.: The occurrence of convulsions in dementia praecox, manic-depressive 

insanity and the allied groups. State Hosp. Bulletin, N. Y., 1914. 

(20) Kirby, G. H.: Catatonic syndrome and its relation to manic-depressive insanity. 

Journ. of Nerv. and Ment. Dis., 1915, lx. 

(21) Rosanoff, A. J.: Intellectual efficiency in relation to insanity. Amer. Journ. of 

Insanity, 1916, lxxiii. 

(22) Rosanoff, A. J.: Survey of mental disorders in Nassau County, N. Y. Report 

published by the National Committee for Mental Hygiene, N. Y., 1917. 


INDIVIDUAL AND RACIAL INHERITANCE OF MUSICAL TRAITS 


CARL E. SEASHORE 

Division of Anthropology and Psychology , National Research Council, Washington, D, C. 

Instead of reporting a particular experiment, I embrace the opportunity 
to lay before this Congress the general result of a number of series of 
experiments. This result or issue may be summed up in the statement that 
musical talent is resolvable into a number of inborn natural capacities which 
may be isolated and measured or rated adequately for statistical or experi¬ 
mental purposes. 

Through a sustained teamwork of collaborators for many years, we have 
gradually designed and made available the necessary measuring instruments, 
standardized procedure, and established norms and other matters of tech¬ 
nique for many of these factors. 

I wish here especially to call attention to their availability in the study 
of racial differences as well as the study of individual differences in the 
experimental investigation of the inheritance of musical talent. The 
relation of these to eugenics is self-evident. 

The approach to the problem of inheritance of musical talent, from the 
point of view of eugenics, divides itself naturally into five stages or tasks: 
(1) the analysis of what constitutes musical talent and the isolation of 
measurable factors; (2) the development and standardization of methods of 
measurement and rating of each of these talents under control; (3) the 
actual field work of measuring sufficiently large numbers of generations in 
selected family groups; (4) the interpretation of such results in terms of 
biological principles of heritable factors; and (5) interpretation and dissemi¬ 
nation of established information for eugenic guidance. 

The first and second of these stages I have treated at length in an in¬ 
troductory way in a recent volume entitled “The Psychology of Musical 
Talent. ” L In attempting such isolation and control of the factors for the 
purpose of measurement and rating we are confronted with numerous 
difficulties, complexities, and dangers, of which frank recognition is the 
first step in scientific procedure. Some of these I have outlined in a recent 

1 Silver, Burdett and Company, 1919. 


231 


232 


CARL E. SEASHORE 


article in The Musical Quarterly 2 on “The Inheritance of Musical 
Talent,” pointing out that musical talent is not one but a group of 
hierarchies of talent, each highly organized; that measurement and ap¬ 
praisal of musical talent must take cognizance of the possession of capacity 
rather than of evidence of achievement; that, although the musical geniuses 
and musical defectives present interesting types for investigation, the 
musical mind is the normal mind; that we must distinguish between the 
talented person and the genius in music; that the versatility and the 
plasticity of the human organism offer most extraordinary resources for 
adjustment under handicap; that scientific procedure encounters peculiar 
difficulty in dealing with the artistic attitude of the musician; that we can 
not appeal to any generally recognized biological theory of mental inheri¬ 
tance; that we must discard the literature on musical inheritance now extant, 
because it is not based on scientific conceptions of the musical mind; and 
that we must, with patience and forethought employ the methods of 
experimental psychology for the analysis of musical talent and for the 
measurement of talents for statistical purposes, in spite of the slowness of 
the procedure. 

On the third stage we have driven an entering wedge in the survey of 
six prominent musical families on which Dr. Stanton is scheduled to report 
at this session. 

The eugenist might rightly expect me to recite established facts on the 
inheritance of musical talent and present arguments showing that they 
should be applied. But the time is not yet ripe for either. The object 
of this paper is merely to present a point of view, showing that such facts 
can be gathered; and this is done in the anticipation that, once established, 
the desirability of their application will be taken for granted by those who 
are interested in this phase of eugenics. 

In the few minutes available for this report, I shall limit myself to some 
remarks on the selection of measurable factors and the organization of 
experiments covering a sufficient number of generations and families. 

We must at once abandon the idea that a person is either musical or not 
musical, that the gift of music is one gift or one talent. In view of this I 
have drawn up in tabular form what, at our present stage of experimental 
analysis, would seem to be isolable factors of musical talent that may be 
inherited. 


2 The Musical Quarterly, Schirmer Company, October, 1920. 


INHERITANCE OF MUSICAL TRAITS 


233 


Isolable Factors of Musical Talent that may be Inherited 

I. SENSORY CAPACITIES 

A. Pitch 

1. The sense of pitch 

2. The sense of timbre 

3. The sense of consonance 

B. Time 

4. The sense of time 

5. The sense of rhythm 

C. Intensity 

6. The sense of intensity 

7. Acuity throughout the pitch range 

H. MOTOR CAPACITIES 

A. Pitch 

8. Instrumental control of pitch 

9. Singing of (1) key, and (2) interval 

10. Rating of (1) timbre and (2) range of voice 

B. Time 

11. Free timed action 

12. Regulated action 

13. Rhythmic action 

C. Intensity 

14. Instrumental control of intensity 

15. Volume of voice 

m. REPRESENTATIVE CAPACITIES 

A. Imagery 

16. Auditory 

17. Motor 

18. Visual 

B. Memory 

19. Auditory memory span 

20. Visual-motor association 

21. Auditory-motor association 

22. Speed and accuracy of learning 

C. Imagination 

23. Imagination type 

24. Creative imagination 


234 


CARL E. SEASHORE 


IV. GENERAL CAPACITIES 

25. Intelligence quotient 

26. Emotional type 

Fundamentally there are three groups of talent corresponding to the 
three elements, pitch, time, and intensity in music; and these are represented 
at four levels, namely, the sensory, the motor, the representative, and the 
higher cognitive and the emotional. 

Sensory capacity for pitch is represented typically in measurements of 
the sense of pitch, the sense of timbre, and the sense of consonance, dealing 
respectively with a simple tone, a rich tone, and combinations of tone as 
in harmony. These are probably closely related and may be regarded 
as one factor dependent upon the character of the tone analyzing mechanism 
in the organ of Corti. Measurements of the sense of time and the tem¬ 
poral aspect of the measurement of the sense of rhythm probably represent 
another entirely independent factor; but we must remember that rhythm 
is a complex of several factors. The sense of intensity measures the nat¬ 
ural capacity for hearing distinctions in the loudness of sound. This seems 
to be quite unrelated to the acuity of hearing, as the sense of intensity 
may remain unimpaired for audible sounds where the acuity is markedly 
defective. 

The motor capacity for the control of pitch depends upon two factors, 
namely, general motor control and capacity for hearing pitch. This talent 
differs in control of pitch in instruments and in voice, each of which may 
be measured separately; and both for instrumental and vocal control of 
pitch there may be distinct and unrelated factors, such as the gift of wide 
range of pitch, or a beautiful tone quality as represented by timbre in the 
natural voice. On the motor side the free timed action and regulated action 
as they appear in the temporal aspect of rhythmic action probably represent 
a single factor. In the motor capacity for control of intensity we may 
recognize the natural capacity for volume of voice and a capacity for pre¬ 
cision in the control of intensity of sound either by voice or instrument in 
musical expression. 

Turning to the capacities for mental representation of music, we recog¬ 
nize distinct variables in auditory, motor, and visual imagery; and, in 
musical memory, we recognize as distinct factors, the auditory memory 
span, visual-motor association as in sight reading, and auditory-motor 
association as in playing by ear. These three taken together are the chief 
determinants of speed and accuracy in the acquisition of skill in music. It 
seems probable that fairly distinct types of imagination may be identified; 


INHERITANCE OF MUSICAL TRAITS 


235 


and capacity for creative work is a comparatively rare gift which may be 
recognized in different types. 

Two general capacities have distinct value in the rating or valuation of 
musical talent, namely, the intelligence quotient and the emotional type. 

In judging the merits of this or any similar analysis of measurable or 
ratable capacities, we must bear in mind that the analysis is made for the 
purpose of controlling conditions in experiment and does not rest upon any 
assumption to the effect that these factors operate singly or in isolation 
in the exercise of music. The isolation of factors of this kind is, however, 
absolutely essential in order that we may deal with specific, repeatable, 
measurable and describable situations. To illustrate, the sense of rhythm, 
auditory imagery, and intelligence are isolable for the purpose of measure¬ 
ment, although in musical appreciation and expression they are integrated 
into one factor. 

However inadequate this preliminary analysis, it paves the way for the 
asking of specific questions, such as, is the sense of pitch, is the sense of time, 
is auditory imagery, is musical imagination, is a voice of large volume, is a 
voice possessing a given tone quality, heritable? 

This list of factors must be considered as tentative and lacking in detail. 
It is merely intended to point the way. Refinement and relationship 
must be based upon further experiment. 

As to the reference of these elements of musical talent to biological 
factors, determiners, or carriers in the mechanism, we are only in a position 
to assert that it may be feasible. 

There should be no confusion from the fact that some of these factors 
are designated from the mental and others from the physical point of view; 
from the psychological point of view, all are psycho-physical; and, in obser¬ 
vation and measurement, we regard a factor for one purpose from the mental 
and for another from the physical point of view. 

Likewise, we must recognize that a given capacity which is an essential 
factor in musical talent may be of general or specific significance in countless 
other mental activities. Indeed, all the capacities here listed have a wider 
scope than for music. Thus general intelligence is not an exclusive musical 
capacity, yet that factor determines the degree of intelligence that may be 
exhibited in the musical talent of an individual. 

Let us turn then for a moment to the problem of the organization of this 
experiment. Granting that we know what we wish to measure and how to 
measure it, the staging of an experiment to determine the laws of the in¬ 
heritance of musical talent in human beings is not different from the 
staging of an experiment on the inheritance of color in peas, the milk- 


236 


CARL E. SEASHORE 


producing quality in cows, or speed in race horses, except that there are 
more factors to take into account and the experiment will take longer. 

There are several ways in which the experiment may be organized. With 
plants and animals you select a colony and breed successive generations 
under observation. With human beings you could select a sufficient number 
from volunteers in which the factor under control is mated in a known way 
and examine them and their children’s children and their mates in successive 
generations. We might, for example, select one hundred newly married 
couples from volunteers, in which we have as large a variety of matings 
as possible with reference to the presence or absence of musical talent, with 
the understanding that an endowment provides for the measurement of 
their children and their children’s mates in successive generations. In this 
way we should follow the very principles that are followed in plant and 
animal colonies for the study of inheritance. There is nothing offensive or 
forbidding in applying this method to musical inheritance in human beings. 
Indeed, the experiment should be most delightful and should present a great 
variety of points of interest for science, art and society. 

Such an experiment would require forethought and stability of organiza¬ 
tion; but it would be relatively inexpensive, since the scope might be limited 
to the number of persons that could be measured by one expert permanently 
employed. Nor would the results be discouragingly remote, because they 
would have cumulative significance from the time of the measurement of 
the first generation, and the remoteness might be shortened by selecting the 
parents at the time the children are of measurable age instead of the time of 
marriage, providing the selection is not made at all on basis of the showing 
of the child. 

In plant and animal experimentation it is customary to take only one factor 
under control and run that intensively, but, in view of the complexity of the 
situation in musical talent, the expense involved, and the necessity for 
carrying the experiment through several generations of human beings 
it would be advantageous to select a large number of measurements, each 
representing more or less unrelated factors, and make these records on the 
members of each generation. Such a list might be selected from the list 
of twenty-six items mentioned in our inventory, or at least on the principle 
of that inventory. 

A more direct approach is to do as Dr. Stanton has done,—select a musical 
individual who is distinguished for his achievement and measure his mate, 
ancestors and progeny, as far as available. This has the advantage of 
grouping about a conspicuously talented person and obtaining significant 
results from a few cases. 


INHERITANCE OF MUSICAL TRAITS 


237 


Another method would be to take selected families who have children of 
measurable age and measure the parents and children at the same time, 
taking large numbers of cases so as to get statistics from matings that repre¬ 
sent great variety of talented and untalented persons. 

It would be bold speculation to try to guess the countless and far reach¬ 
ing scientific facts on the inheritance of these mental traits that might be 
discovered by such methods of procedure. The experiment is particularly 
worth while because musical talent lends itself well to measurement and 
the possession of this gift is of great human interest. 

It is a matter of common observation that we make fairly definite pre¬ 
diction of the chance of reproducing a given quality, such as shortness 
of stature, and talk wisely about the chances of this when two short people 
mate, when one short and one tall mate, when two short people all of whose 
ancestors on both sides were short, or one whose ancestors on one side were 
short and on the other side tall, etc., in various combinations in receding 
generations. And from the experimental work that has been done on 
plants and animals, we are finding the explanation of these common sense 
predictions and establish greater confidence in the predictions. The same 
is, to some extent, true about the inheritance of musical factors. Suppose 
that we substitute for stature, the sense of pitch, the sense of rhythm, 
musical memory, musical intellect, creative imagination in music, register 
of voice, range of voice, volume of voice, etc., and regard these as independent 
factors. Our common sense observation gives us some feeling of assurance 
in the power to predict these; and it is the aim of this paper to maintain that 
a scientific knowledge of heredity will increase our power of prediction of 
musical talent in the same sense that it has advanced the prediction of 
stature. 

Is such knowledge worth while? Is the rose less beautiful to the botanist 
than to the ignoramus? Is the choice of a mate less of a love affair because 
one has the power to understand and know some of the laws of life 
of the thing he adores? When the first baby came into my home there was 
a “ladies’ aid” meeting across the street, and it is reported that one of the good 
ladies exclaimed, “ That poor Seashore child! ” All ears were open for alarm. 
“What is the matter?” The reply came, “His father is a psychologist!” 
Yet I have a feeling that my love for my children is none the less genuine 
and intense for the fact that I have some scientific knowledge of child life. 
Should not my love for my children as compared with the love of one who 
knows nothing about child nature be as the botanist’s or horticulturist’s 
admiration for a flower is as compared with the ignorant peasant’s admira¬ 
tion of the same flower? 


238 


CARL E. SEASHORE 


In other words, suppose a youth is endowed with a wonderful musical 
mind and imbued with a deep desire to have children endowed with the 
same precious gift. Would it be an advantage to have definite knowledge 
and insight in the predictions on the basis of family history, for each of the 
factors I have described, such as, the sense of pitch, musical imagination, 
and quality of voice and be able to say that the chances are 10 to 1, 1 to 1, 
1 to 2,1 to 5,1 to 10, or 1 to 100 that this factor shall be reproduced? 
If youth enters into wedlock in possession of this knowledge, will that not 
tend to a deepening of love to the extent that it carries with it confidence of 
great promise? The opponents of eugenics fail to realize that scientific 
facts when well established become a part of common sense and a basis of 
intuitive reactions. My proposition is that if certain musical talents are 
heritable, as we believe them to be, it is quite within the power of future 
generations to enhance the quality and degree of a musical talent by con¬ 
scious selection. The great significance for eugenics, however, lies not in 
the development of a system of artificial and formal eugenic guidance, but 
rather in the popular assimilation of well established facts in the common 
sense of the age and the naive projection of this common sense in natural 
reaction in courtship and mating. 


AN EXPERIMENTAL INVESTIGATION OF MUSICAL 
INHERITANCE 

HAZEL M. STANTON 
State University of Iowa 

This paper is a report of a recent investigation of the inheritance of 
musical talents made from the Eugenics Record Office of the Carnegie 
Institution of Washington. The purpose of initiating such an investiga¬ 
tion was to establish technique for studying the inheritance of musical 
talents by means of quantitative measurements. Four of the Seashore 
measures of musical talent, viz.: The sense of pitch, the sense of intensity, 
the sense of time, and tonal memory, were given to members of families in 
which one member was known to be conspicuously talented in music. 
These measurements were developed and standardized from the Music 
Laboratory Studio of the State University of Iowa under the direction of 
Professor C. E. Seashore, head of the department of philosophy and psy¬ 
chology. During the past eighteen years extensive experimentation has 
revealed the apparent basic nature of these measurements and has shown 
them to be little affected by practice, age, musical training, sex and intelli¬ 
gence. By this I mean that when a reliable result is obtained from an 
individual, for example, a superior record in the sense of pitch, there will 
be little or no variation in this record on repetition of the measurement under 
similar conditions, children may do as well, better or poorer than adults, 
the same measure may be given to those with no musical training as to those 
with musical training. We are measuring basic capacities, capacities which 
a child possesses by right of birth and not what he acquires during training. 

The measurements of intensity discrimination, time discrimination and 
tonal memory were given by means of phonographic records on which stimuli 
from the standard laboratory apparatus have been recorded by Professor 
Seashore. For the measurement of the sense of pitch in terms of pitch 
discrimination, the standard pitch discrimination forks with resonators 
were used in place of the phonographic record, first, because the forks are 
better adapted for very fine measurements, and second, in order to intro¬ 
duce variety in the procedure. 

Pitch discrimination was measured by pitch intervals ranging from 30 
d.v. to | d.v. in a tonal region of 435 d.v. The observer discriminates 

239 


240 


HAZEL M. STANTON 


between two tones differing in pitch, the second tone presented being 
higher or lower than the first. For the purpose of intensive work the in¬ 
dividual method of constant stimuli was used. The threshold values, 
computed from conversion tables, may be determined to tenths and hun¬ 
dredths of one vibration. 

Intensity discrimination, time discrimination and tonal memory were 
studied by means of the serial stimuli method. Intensity discrimination 
is a measurement of the capacity for discriminating differences in intensity 
or loudness. This measurement contains five graduated steps, the first 
easily perceptible, the last difficult to perceive. 

The measurement of time discrimination refers to an individual’s capac¬ 
ity for discrimination between two time intervals, the second interval 
presented being longer or shorter than the first. The stimuli cover a range 
of steps from an easily perceptible difference of 0.20 of a second to a differ¬ 
ence of 0.02 of a second. Such a measure of an individual’s capacity for 
discriminating time intervals is not a measure of rhythmic perception but a 
measure of one of the basic constituents of rhythm. It gives one partial 
knowledge of the sensory aspects of rhythm, the receiving of an elementary 
impression of time. 

The measurement of tonal memory is a measure of immediate memory 
for a span of tones. It consists of five steps gradually increasing in difficulty, 
each step containing a certain number of successively presented tones, 
followed by a second span of the same tones with the exception of one tone 
which is changed in pitch. The observer identifies the changed tone by 
indicating its number in the group. The spans increase gradually in 
presentation from a group of two tones to a group of six tones. 

These four measurements were supplemented by a special interrogation 
dealing with individual case histories, individual musical experiences, 
including musical environment in the parental home and in the community, 
musical education and training, musical activity, musical appreciation, 
musical memory and imagination, family musical history, and a short asso¬ 
ciation test. 

In selecting the musicians from whom a family study could be developed 
we chose those who were available for an interview during the months re¬ 
served for their section of the country and who had families, the members of 
which were significant in number and available for appointments. In so 
far as possible a family study included all the members of the restricted 
family. Individual interviews averaged two hours in time. The four 
quantitative measurements were given to 85 individuals comprising six 
families. The qualitative information was obtained for 531 individuals. 


INVESTIGATION OF MUSICAL INHERITANCE 


241 


# The order of presenting the measurements and the supplementary informa¬ 
tion to an individual was, first, the measurement of pitch discrimination 
with the tuning forks; second, the free association test; third, the three 
measurements on the phonographic records, viz., the sense of intensity, 
the sense of time, and tonal memory; fourth, the systematized interrogation 
covering case histories and musical experiences. 

Norms for each measurement have been established for three age groups, 
viz., fifth grade pupils, eighth grade pupils, and adults. 

The results of the quantitative measurements are presented as follows: 
the final score of all the measurements is determined in terms of percentile 
ranks consisting of a standardized distribution of results ranging from 1 to 
100. The individual ranks are presented in the form of individual talent 
charts as seen on the pedigree talent chart below. One family is charted 



with the individual talent profiles for every member to whom the measure¬ 
ments were given. The percentile rank obtained in each measurement is 
presented in the horizontal sections. The sense of pitch rank is recorded in 
the upper section, the sense of intensity in the second section, the sense 
of time in the third section, and tonal memory in the lower section. All 
ranks extending to the right of the middle line of each chart are above 
average, those to the left are below average. Time does not permit dis¬ 
cussion of these charts in their relation to musical experiences for each 
individual. 

The results of the qualitative information have been analyzed and rated 
into three groups of A, indicating high rating; C, average; and E, low. 
By direct comparison of the results obtained in the measurements and the 
qualitative ratings of musical information, those possessing the highest 5 
per cent talent profiles are also high in ratings of musical experiences; those 
possessing the lowest 5 per cent talent profiles lack musical interest, and have 
experienced no musical activity. 







































































































































































242 


HAZEL M. STANTON 


The results of the measurements when distributed according to the six 
types of matings, viz., superior x superior, superior x average, average x 
average, superior x poor, average x poor, and poor x poor, show the follow¬ 
ing distribution of offspring for each measurement in so far as examples of 
each type were obtained. In pitch discrimination both parents superior, 
of 16 offspring 15 were superior and 1 average; one parent superior and 
the other average, of 11 offspring all were superior. 

In intensity discrimination, both parents superior, of 7 offspring 6 were 
superior and 1 poor; one parent superior and the other average, of 6 
offspring 5 were superior and 1 average; one parent superior and the 
other poor, of 11 offspring 5 were superior, 4 average and 2 poor. 

In time discrimination, both parents superior, of 5 offspring all were 
superior; one parent superior and the other average, of 6 offspring 3 were 
superior, 2 average and 1 poor; one parent superior and the other poor, 
of 8 offspring, 5 were average and 3 poor. 

In tonal memory, both parents superior, of 5 offspring 4 were superior 
and 1 average; one parent superior and the other average, of 17 offspring 
14 were superior, 2 average and 1 poor. 

In partial answer to the question regarding the tendency of musical or 
non-musical parents, of musical or non-musical stock, to have musical or 
non-musical children, the following statements based upon 22 matings 
have been deduced from the qualitative data. Of 11 children of musical 
stock all were musical except 1 whose normal growth was stunted. Of 
25 children of non-musical stock all were non-musical. Of 17 children 
from parents, one of whom was musical and the other non-musical, 6 
children were very musical and 11 were non-musical. Representative mat¬ 
ings of the other types were insignificant in number. 

In presenting this study to you I call your attention to a few of the out¬ 
standing difficulties and limitations immediately encountered; first, the 
limited number of individuals upon whom preliminary conclusions are 
based; second, the accumulation of results into the average and above 
average groups, leaving a paucity of results in the poor group; third, 
the extreme variation in age ranging from eight years to eighty years, 
which adds new problems to the experimental phase of this and further 
investigation. 

This approach, however, gives significant promise toward the establish¬ 
ment of definite laws of the heredity of musical talent by determining 
the type of musical sensitivity of every member of the family whether or 
not an opportunity for musical expression was utilized. The value of in¬ 
formation obtained from technique of this kind may be foreseen in its possi¬ 
bilities of eugenical application and prediction. 


THE CONSEQUENCE OF WAR AND THE BIRTH RATE IN 

FRANCE 


LUCIEN MARCH 
La Societe franqaise d’Eugenique 

As a result of the War, the France of 1914 has lost 1,400,000 of her in¬ 
habitants in the prime of life, most of them fit for producing children. On 
the other hand, among the survivors of the fighters of the Great War, a 
certain part of the 800,000 total invalids will never be able to give birth to 
strong healthy children, either because they are no longer capable of marry¬ 
ing, or because they are affected with tuberculosis or other constitutional 
maladies. 

To these direct losses must be added the loss of births. Before the War, 
the number of living births balanced with a slight excess over the number 
of deaths; the annual number was about 750,000. During the six years 
1914 to 1919 inclusive, the deficit, in comparison with six times this mean 
number, reached 400,000 births which ought to have survived normally, 
and which were lost owing to the War. 

On the other hand, deaths in the civil population have been more numer¬ 
ous than formerly, so that 400,000 more deaths are added to the 1,400,000 
missing births and to the 1,400,000 soldiers killed in war, giving a total of 
more than 2,000,000, taking account of possible useless counting and of 
immigration. These results are calculated besides in supposing that, in the 
invaded regions, the loss, estimated proportionally to the number of in¬ 
habitants, be the same as in the uninvaded territory; on the other hand, 
the numbers are applied to the territory of 1914, but Alsace and Lorraine 
cannot nearly fill the loss of population of this region. The provisional 
results of the census of 1921 confirm these suppositions. 

But that is not all. Privations have broken down the health of many 
children born during the war or a few years before, especially in the regions 
of the Northeast, where, during the German occupation, they have lived 
in a state of veritable physical misery. Indeed, infant mortality, even in 
the uninvaded districts, has been notably higher during the war than be¬ 
fore, in spite of the low birth rate. 


243 


244 


LUCIEN MARCH 


Finally, a certain increase of alcoholism, tuberculosis, venereal disease 
and various nervous diseases may influence unfavorably the vitality of the 
nation and the race. 

Many years will be necessary to repair the losses of population, direct or 
indirect, chargeable to war or to the evils which have accompanied it. 

To avoid the unfortunate consequences of these miseries, certain people 
believe it is necessary to induce procreation by all possible means, and do 
not fear an excess of population for a long time. 

Others think it proper that each man of proper age to have children have 
the three or four children necessary to permit a moderate increase of popula¬ 
tion. Others believe that a continued increase of population would create 
an economic peril and contain the germ of future wars. 

Others wish certain restrictions, above all in the poorest of the population, 
to improve the quality of this population. 

The considerations which are the most important are the following which 
shall be examined from the point of view of Eugenics and the point of view 
of Economics. 


i 

Today, respect of human life in all its degrees makes us condemn in¬ 
fanticide and abortion. There only remains, then as a means of artificial 
selection, the prevention of births. 

But the universal concern which determines parents to limit the number 
of the children is the burden, at least momentarily, which the latter 
represent. 

Also the question of the birth rate, in its entirety, and save some excep¬ 
tion of which we shall speak soon, comes back again to a question of 
economic morale. For physical passion finds play without producing the 
being which is its end, and this being is often today the payment of a 
sacrifice freely agreed upon. 

Humanity ought not to perish by its own error, any more than the col¬ 
lections which form the cells of it. Such is the higher principle which ought 
to reconcile itself with the practical impossibility of multiplication without 
limit. 

According to etymology and the definition given by Gal ton, eugenics is a 
general study of the improvements of which the race is susceptible, race 
being characterized by common physical or mental qualities which mani¬ 
fest themselves in certain groups of men and differentiate them from other 
groups. Two ideas enter here, that of improvement, that of race. To 
what realities do they correspond? 


THE WAR AND THE BIRTH RATE IN PRANCE 


245 


We cannot define progress, the process of making perfect, but, when we 
look back, we feel the differences which separate the life of other times and 
of the present; evolution appears to us to follow a certain direction. We 
can then legitimately propose to continue life in this direction. 

In the second place, although in a biological sense pure human races are 
not numerous, one can prove that a number of groups of individuals are 
-distinguished by their physical and mental characters, apparent and dis¬ 
tinct as a whole, from another group. Without modifying these characters 
to the point of making the differences disappear, one can improve their 
manifestations, the manner in which they act in each human group, that is 
the aim which Eugenics seeks. But we must not lose sight—for other 
sciences seek the same end, the science of education for example—that 
Eugenics is occupied, it seems, only with the measures capable of effects 
upon descendants, that is to say, transmissible by heredity or capable of 
operating a selection advantageous for future generations. 

This science has not yet its general principles well established. It is not 
yet settled, it is still in a period of growth. And that permits some liberty, 
some differences of viewpoint to those who try to improve it. 

There are however acquired facts, indisputable connections; for the pres¬ 
ent one may withdraw to this ground. 

Whatever may be the opinion as to the relative importance of one or the 
other factor, heredity or environment—that is the principal point on which 
personal opinions oppose each other—one cannot deny the influence of 
heredity. Physical and mental resemblances of parents and children are 
very evident; the hereditary transmission, at least in the most closely re¬ 
lated generations, of certain physical peculiarites, such as stature, conforma¬ 
tion of the skull, hemophilia, polydactylism, etc., or of mental defects 
such as epilepsy, certain forms of mental deficiency or feeblemindedness, 
are today almost proven. Provided that it is always a question of the 
simple characters whose existence can be removed. Resemblances between 
children born of the same parents do not prevent sometimes great differences 
appearing in these children. The heredity of abilities or that of defects 
has nothing of fate: then education may modify nature. 

As to the influence of environment on the mode of development of the 
created being, whatever may be its importance for this being itself, the 
question which interests eugenics is to know if this influence acts upon the 
descendants after being hidden a number of generations. On this point, 
certain savants, Weissman in particular, have declared negatively. Others 
have shown, by experiments on inferior organisms, that organic modifica¬ 
tions performed on these organisms are transmitted to their descendants. 


246 


LUCIEN MARCH 


As Dr. Apert has remarked in France, as far as man is concerned, it seems 
that only the modifications relating to the nervous system have yielded, up 
to the present, observations truly conclusive. Yet the interpretation of 
these facts has been contested; they have been charged to hereditary pre¬ 
dispositions, but it is always easy to draw into the results of an observation 
the effect of a hidden influence as mysterious as that of heredity. 

One does not know enough to challenge on these obscure questions, and 
yet of such great importance to humanity is a development, sustained and 
growing, of scientific researches relative to the heredity of man that this is 
the desire of all those who are interested in eugenics. 

The action of the two factors of the transmission of character, from one 
generation to another, works by means of germplasm, but this is guided by 
selection: natural selection by death, artificial selection by sexual union. 

M. Edmond Perrier, the very regretted president of the Societe frangaise 
d’ Eugenique, gave recently an opinion, that, in primitive nature, natural 
selection had not perhaps had the exclusive effect which the Darwinians 
have attributed to it. 

Moreover, what precisely is natural selection? Does one understand 
simply that an individual incapable of adapting himself to the conditions 
imposed on him by environment disappears and they only survive who are 
capable of adapting themselves? That does not add a great deal to our 
knowledge, as M. Balfour (speaking before the First International Eugen¬ 
ics Congress) remarked, since that amounts to saying that only those are 
capable of surviving who survive, a veritable truism. 

And if one means that only those survive who are capable of surviving, 
M. Perrier answers (Eugenique, mai 1921, page 197) that those who are 
incapable of surviving in one determined point can escape death by flight, 
and it is thus perhaps that at first the living world has evolved. 

In truth, death and survival are a form of selection from which may re¬ 
sult for humanity, as for every society of living beings, good or evil following 
the individual in the case, and surrounding circumstances. 

If we are unable to modify the innate qualities of the individual, we may 
often, by acting upon the circumstances which surround them, make useful 
the qualities which they have. 

This is one of the essential duties of eugenics: to favor and encourage 
the work of health and the work of education of promoters of social progress. 

As to artificial selection, we may endeavor that the revival of man by 
birth shall be facilitated for those who possess the best qualities, be shackled 
for those who show, on the contrary, defects and faults. However, we ought 
to ask ourselves whether there does not exist now and then a certain 


THE WAR AND THE BIRTH RATE IN FRANCE 


247 


opposition to these two directions: that which leads to the improvement of 
conditions of existence and that which leads to the research of the best 
qualities in the descendants. 

Opposition has been noted many times, especially among the English 
eugenicists. Nature, they say, in a convenient anthropomorphic language, 
nature has arranged itself for the beings least endowed for life discourages 
before those who are better endowed. This observation is just; in admitting 
that as to the shadowy origin of life, flight has been a means of preservation, 
this means is not worth much when it is impossible to flee from danger. 
This is the case when illnesses and bodily struggles have made disappear the 
least worthy beings, the least capable of resistance. But when human 
fraternity, pity, science, and hygiene unite their efforts to defend the weak, 
many individuals who would have disappeared, left to themselves, live in 
spite of their inabilities and transmit these to their descendants. As is 
often remarked, the humanitarian tendencies of our time, our social legisla¬ 
tion and all the measures which come from the same principle, have this 
effect—of which people are not sufficiently warned—to oppose the play of 
natural selection. This manner of thinking bears a great deal of truth. 
However, no defender of eugenics thinks of suppressing pity, or hygiene, 
to reestablish natural selection in its barbarian despotism. The efforts of 
humanity tends to utilize the natural forces for their own ends and not to 
let them act blindly. Also when the ideal of healthfulness and social 
progress is opposed to the ideal of perfection of race, because the first is 
contrary to the effect of natural selection, it becomes necessary to demand 
from artificial selection much more important effects, and especially 
those better regulated, than those which she produces among primitive 
people. 

It is this which we will now take account of in passing to the special 
question of birth. Even though we can lessen the effects of natural selec¬ 
tion, we can much more surely intervene by artificial selection to favor the 
perfecting of the race and above all to prevent its degeneration. The 
point is to make good use of this power. 

ii 

In all times man has tried to deal with the multiplication of his race. 
Independently of wars, famines, epidemics, whose destructive effects extend 
themselves over entire populations, suppression of infants already born, 
abortion, and prevention of births have been practiced. 

Eugenics, as well as economics, can tell us what the social interest 
demands, to be sure. From the point of view of eugenics, the experience 


248 


LUCIEN MARCH 


of centuries and of many researches teaches us above all that there are 
transmissible defects, multiplications of which must be avoided at all costs. 
These are notably the hereditary predispositions to insanity, to mental 
weakness, to epilepsy, and to detrimental malformations; or again the ac¬ 
quired dispositions chargeable to the poisons of the nervous system, such 
as alcohol and the spirochete of syphilis. 

Evidently one cannot always be sure in advance of the effect of those 
influences which, acting in the mass, give rise to differences. Nevertheless 
there are individuals whose duty it is not to procreate, not to give birth to 
beings whose chances of deformity or mental deficiency are really too great. 
This duty is all the clearer when one is forced to conserve the life of those 
beings who, in other times, would have been condemned to a more rapid 
death by the brutalities of existence. 

Beside the circumstances which justify and command abstinence, there 
are still others which can be drawn in very legitimately to limit the number 
of children; one understands, for instance, that, in the very crowded urban 
districts, the smallness of homes, the promiscuity, cause an excessive mor¬ 
tality when the families are large, without having means for choosing spa¬ 
cious homes. Finally, there are individual proprieties worthy of respect, for 
example, the care of the mother’s health when she cannot stand numerous 
pregnancies, not to speak of the limits which can impose a legitimate fear 
of an undeserved loss, if a large family imposes a burden which surpasses 
the strength of this family. 

One would not then accept the formula of an unfortunate equality, which 
would impose on all adults the obligation of having a determined number of 
children, any more than one would dream of recommending an unlimited 
fecundity. It is necessary to discard these precise but too simple formulas, 
and to keep within the bounds of asking that each adult of age to have 
children have them if he reasonably can. Each one, in fact, has the duty 
of transmitting the life that he has received, and even of improving the 
value of that life as those who have preceded have striven. And thus is 
imposed, according to the limits of his means and his capacities, the duty 
of perpetuating the family to which he belongs, the duty of contributing 
to the aim of his country and the progress of all humanity. 

The formula is doubtless very vague, it is addressed to conscience, for it 
is conscience alone which is the judge of the degree to which the order has 
been obeyed. It is the same as when one appeals to the conscience of each 
one to participate in the defence of country or of national burdens. 

In this case, it is true that legislation adds itself to the moral obligation: 
is it not necessary that legislation also intervene in favor of the birth rate? 


THE WAR AND THE BIRTH RATE IN FRANCE 


249 


The answer to this question is not doubtful; we cannot omit a certain social 
organization capable of stimulating conscience and assuring the desired 
result: that is to say, the number of births which appear necessary for the 
whole of the population. 

However, two objections have been made. One declares that before 
increasing the birth rate it would be better to reduce mortality and, above 
all, infant mortality. 

It is obvious that all measures capable of reducing mortality are good in 
themselves. But, since the remotest historical times, it has not appeared 
possible to lengthen the maximum of human life. We can only hope to 
lengthen the mean duration of life. But that will not produce an appre¬ 
ciable increase in the population in the countries where the number of births 
depends on familial foresight, when the parents determine, so to speak, in 
advance the number of children they will raise. Three years out of four in 
France, the number of births in one year is related to the number of infants 
who have died in the preceding. If many children die they are replaced, 
if there are not premature births, so many births less. 

The second objection is that instead of seeking the number of children, 
it is preferable to concern oneself about the quality. We have seen that the 
quality of population is in fact the principal aim of eugenics. 

But it is proper to consider successively the family and the nation. 

In the family, at the time when the number of children does not exceed 
the reasonable limit of which we have spoken, one can affirm that quality 
goes hand in hand with quantity, far from being opposed to it. One has 
often noted the case of the only child. One has also cited examples of num¬ 
erous brilliant men who are among the younger members of families, some¬ 
times of very high rank. 

As to the Nation, she may claim a certain choice, a selection, of which we 
have mentioned the importance in the first part of this paper. 

But by what sign may inferiority and superiority of qualities be recognized, 
it being understood that one is to be forced to prevent from procreation the 
bearers of defects?' It is proposed to take wealth for an index. Numerous 
inquiries have proven in fact that in the slums of cities, among the individuals 
who have no care for the morrow, are found the greatest number of trans¬ 
missible defects and the most afflicted children. On the other hand, mani¬ 
festations of intelligence and various abilities have appeared more frequently 
in the children of well-to-do families than among those of poor families. 

But the influence of environment is considerable here, even as that of 
education. If one leaves out the small part of the population which is 
composed principally of social outcasts, one cannot but affirm that the innate 


250 


LUCIEN MARCH 


qualities (we do not speak of acquired qualities) are less in the families of 
small income than in the families of large income, especially if one takes 
into consideration all classes of population, city and country, intellectual 
and artisan. 

Reserving the elimination of undesirables, it does not seem that there is 
serious reason, from the single point of view of eugenics, of seeking births 
in one class of population more than in another. The numerous statements 
which have been made on the retrogression or even the degeneration of 
families which have not renewed themselves sufficiently, tends on the 
contrary to provoke the incessant mixing of social classes rather than their 
separation. When one considers the state of the population, one perceives 
great differences in the birth rate. In France, the birth rate is generally 
greater in the country than in the city, greater in the mountainous regions 
than in the valleys, greater among the agriculturists, the sailors, the fisher¬ 
men, colliers of the North, the heads of great industries, than in the middle 
classes, among the artisans and especially among the clerks. These dif¬ 
ferences explain themselves; they appear in the nature of things, and, for 
the moment at least, they do not carry any danger. We know that de¬ 
population does not reach the towns which are being filled unceasingly 
by an influx of inhabitants from the country; it is then the birth rate in 
the country upon which effort should principally be brought to bear; it is 
there that results can be gained most easily, at the least expense and under 
the best conditions from the point of view of hygiene, as well as from the 
point of view of eugenics. 

Moreover, social action ought not to confine itself to facilitating the birth 
of children; it is still necessary that institutions be permitted to raise 
children up to a certain age. Questions of education, emigration and 
immigration are also questions on which eugenics has something to say, 
especially on the question of immigration which has gained since the war 
an importance and character unknown in France before. 

Eugenics has also something to say on the psychological and moral side 
of the question of birth rate. Prevention of births, regarded as necessary 
in a certain measure, can be recommended as only according to the means 
indicated by Mai thus; the delay of marriage. 

Fecundity of marriage, which one supposes as great as to allow the care 
of maintaining a L healthy family well adapted to life ought not to be 
fettered by an excessive fear of life, or by the fear of effort. All hope of the 
future cannot be realized except with a certain present sacrifice. It is 
necessary to make some personal sacrifices and to have hope in the future. 


THE WAR AND THE BIRTH RATE IN FRANCE 


251 


These sacrifices will be moreover fruitful for posterity. In what measure 
can they be shared; what profit can they yield for it? That is what the exam¬ 
ination of the question from the point of view of economics will show. 

hi 

The economic power of a country depends primarily on its producers: 
that is to say, on those who by their work make serviceable natural riches. 

Now we have seen above what is the loss of population since the war. 
The loss comes principally from the avoidance of marriage. During the 
war, many young men rightly wished to reach the end of hostilities before 
marrying. From that has come the increase of marriages in 1919 and 1920. 
The same phenomenon has been observed after all wars, it is easily 
explained. 

But the deficit is none the less a noteworthy fact in our country and 
in Belgium. While the population of Great Britain has increased by 
1,300,000 during the same time and that of Germany has hardly diminished; 
if it has diminished at all, we are still ignorant of it. We know in any case 
that in Prussia the number of deaths has not fallen below the number of 
births. 

Therefore imagine the state of the French population in fifteen years. 
At that time, there will be lacking, taking account of the mortality, 500,000 
young men of the ages of fifteen to twenty-one years, a loss which must be 
added to the 1,400,000 men of eighteen to fifty years of age killed during the 
war, and who would then be thirty-three to sixty-five years old, as well 
as the 500,000 young men of the same ages who have died in the civil pop¬ 
ulation in excess of the normal mortality. In all, about 2,000,000 individ¬ 
uals would be counted in the male population to fifteen to sixty-five years, 
or about one-sixth of the population. 

There would be lacking in consequence, about 1935, one-sixth of those 
whose work must furnish the principal source of income of the nation. In 
spite of the restoration of Alsace-Lorraine which brings us 400,000 adults 
of fifteen to sixty-five years, but which demands also workers for its fields 
and iron foundries, it is certain that the French production will be deprived 
of an important part of its active forces and that the economic life of the 
country will languish for many years if energetic measures are not taken 
without delay to ward off the threatening deficit. 

Without doubt, one might temporarily make appeal to foreign workers; 
however, assimilable populations can only furnish a small part; it will be 
necessary to have recourse to unassimilable races very different from ours, 
and which will furnish quickly undesirable elements. 


252 


LUCIEN MARCH 


Doubtless also the deficit of masculine workers has caused the more 
general employment of women. But the women who work cannot be fruit¬ 
ful mothers. Feminine work will only be a short-lived mitigation. 

For all time, since the infant demands care and pain, as well as the joys 
he brings, maternity has been a cause of care and effort. Among primitive 
tribes which are malplaced, it is obligatory not only to nourish but even to 
bear these children. In our civilized societies, and especially in urban 
centers, where civilization is* most refined, the burden is often very heavy, 
the difficulties of lodging, the hindrances of traffic, the care for appearance, 
which is applicable to children as well as to parents, the care for the health 
of the mother and all the complications of urban life; the laws for working 
women, the educational obligations and the impossibility, in poor families, 
of using the work of young children, make heavy enough the maintenance 
of even a limited number of children. 

Formerly, in poor families, who are the most numerous, the help which 
grown children gave to their old parents, compensated in some measure 
for the privations which they had caused at first. Today, collective in¬ 
surance is substituted for this kind of family insurance of previous times. 
In consequence, the child usually never brings any repayment in exchange 
for what he cost. Also the care for his future invited the foresigh ted parents 
of our time to assure themselves of the excellent probabilities of his future 
establishment, which determines them also to restrain their responsibility. 
When the children may rather quickly be an aid to the family, the burden 
is much lighter. Also one finds the greatest number of children among the 
people chiefly concerned in agriculture, and, in every country, in the rural 
populations. 

However, the first obstacle to birth is the possibility of raising the 
children. Doubtless this obstacle exists for many animal species and does 
not hinder their fecundity, but in those species there is no reasoning power, 
no foresight, no respect of life, at least in a degree comparable to that which 
may be observed in civilized human society. 

A second obstacle, which does not exist in any degree outside of humanity, 
is the foresight of parents exercised beyond the time of growth of their 
children. It is not all to have brought into the world children and to have 
raised them to an age when they have strength enough to answer for them¬ 
selves; the environment in which they are placed must permit them to 
live. To understand the economic mechanism of the phenomenon of birth 
it is convenient to distinguish three orders of circumstances: 

1. The means of keeping children alive during their growth. 

2. The eventual means by which these children can live by themselves 
after growth. 


THE WAR AND THE BIRTH RATE IN FRANCE 


253 


3. The opinion of parents on these future circumstances. 

It is necessary to understand here by means of life, the means of leading 
a certain kind of life; one can say in general that it is a kind of life at least 
equal to that to which the parents are accustomed. Often even, the desire 
of the parents is that their children will reach a higher stage of life. 

But the means of living are governed in part by circumstances external 
to living beings and in part by the circumstances which depend on these 
beings themselves. The analysis of these circumstances composes what is 
called the theory of population. 

Long before Malthus, who formulated this theory, they had estimated 
the faculty of increasing the human species, a faculty analogous to that of 
every other living species, when no limitation intervenes. It is wrong to 
reproach Malthus with having employed the formula of geometric pro¬ 
gression, since a simple reasoning founded on a not dissimilar hypothesis 
establishes it. Where Malthus appears to be mistaken is when he attempts 
to justify his law by experience or to deduce from one isolated experience 
the reason of progression. If he could have extended his observations still 
farther, he would have seen that this reason was not constant, and indeed, 
in consequence, progression was not geometric. 

If on the contrary one keeps to the domain of hypothesis, as others had 
done before Malthus, if one supposed that nothing limits the fecundity of 
women, as a woman can bring into the world at least 8 children, taking 
account of cases of involuntary sterility and physiological mortality, it is 
easy to understand that in thirty years a population not meeting any 
obstacle would increase itself in the proportion of 1 to 4 at least: that is to 
say, it would be more than doubled in fifteen years. 

Malthus admitted that the population of the United States doubled every 
twenty-five years; a more rapid progression has been cited, that of the 
Hebrews passing through Egypt: 70 adults became 500,000 in two centuries, 
which means a doubling in exactly every fifteen years, and corresponds to 
the period of doubling of capital placed at interest of 5 per cent a year. 
Every one knows what a fantastic sum is reached with a sufficient number 
of periods of doubling. If the doubling every fif teen years had taken place 
since the beginning of historic times, the men living in our time not only 
could not find place on earth, but would even fill the space which separates 
our globe from far distant stars. Therefore the hypothesis which leads to 
an idea of constant geometric progression is not verified by facts. In 
reality the matter changes with the times because of obstacles which meet 
the indefinite multiplication of a species, for men as well as for all hying 
beings. The interest of the work of Malthus is that this author has classified 
the obstacles and made a choice. 


254 


LUCIEN MARCH 


A second error, which is often made, consists in assigning also a general 
law to the development of the means of existence: These can only increase 
by following an arithmetic progression. 

This pretended law has no theoretic foundation, even in admitting that 
one works in a limited territory, since the production of subsistence depends 
on putting to work the means of production. In fact the means of existence 
has progressed much more rapidly in certain epochs than in others. In 
the nineteenth century for example, the population of the most civilized 
states increased more rapidly than during the previous centuries. There is 
then no general law to be assigned to the increase of population. 

If one applies the formula which would recapitulate the theoretic move¬ 
ment of population, one would begin to say that population develops itself 
in the same measure as the means of living are developed, that there is a 
correlation between the two phenomena. But this vague formula is only 
pure tautology, since one cannot conceive of a population which would 
develop without means of life. Such a formula can only serve as armature 
for a true theory of population. In order to have a theory, one must 
indicate some mechanism for the relation between population and the 
means of subsistence. 

The theory of Malthus tends to establish the fact that individuals, 
according to nature, have an action weaker than the reaction exercised by it. 
Inversely, other theorists, before Malthus the mercantilists and popu¬ 
lationists, after Malthus the advocates of patriotic fecundity, have pre¬ 
tended that, in certain limits at least, man could always obtain from Nature 
what he needed to live. These two theories have been translated by 
picturesque formulae. 

Where bread is born, man is born, say those who believe in blind fecundity 
and limited productivity. Where man is born, bread is born, answer those 
who measure the limitation of fecundity and have faith in the powers of 
invention. 

In reality these brief formulae are too general: following the epochs, 
following the countries, natural increase of population tends to diminish 
production; in other cases the contrary is true. 

In China, in India, the population being increased to a certain degree, a 
deficient production results in veritable hecatombs of human beings, after 
which equilibrium is restored. In other countries where patriarchal life 
has given place to a complicated organization founded on the division of 
labor and the specialization of services, the means of production increase 
sometimes to such a point that production surpasses the needs. In this 
case it is true, the conditions of existence of the people are in a mutual 
dependence, and this dependence gives rise to terrible conflicts. 


THE WAR AND THE BIRTH RATE IN FRANCE 


255 


In the human species, as in all living beings, death appears as an inflexible 
regulator of the interaction of the two factors of life: natural fecundity and 
nourishment. But, in the human species, the individuals are capable of 
foreseeing in some measure future events; foresight is the principal instru¬ 
ment of progress of the species and of civilization. Malthus has marked 
well this difference between human species and others, and he has declared 
that for the brutal regulator of other species one may substitute that of 
reason. That has been expressed, in rather rude form, by a German econo¬ 
mist, Julius Wolf, who sees in the universal diminution of the birth rate the 
effect of increasing rationalism of a life. 

Only, Malthus has not seen the importance which this factor will have and 
the danger which will result when this factor is capable of suppressing all 
the principles of life. He believed, on the contrary, that the power of in¬ 
stinct would always be stronger than the fear of overpopulation, and he 
impregnated the thought of his century with a dangerous pessimism. 

But is it true that increase of population is necessarily a menace to the 
existence of this population? The facts answer for themselves. Not only 
has the nineteenth century seen the civilized nations increase in proportions 
unknown in the preceding centuries without these nations having suffered 
want: but, among them, the most rapid increase in wealth has gone with the 
most rapid increase in population. In England at the beginning of the 
nineteenth century, poor laws imposed excessive burdens on the parishes, 
misery ruled and the lamentable state of the population at the beginning of 
the age of machinery justified later, in the eyes of Karl Marx, its attacks 
against the capitalistic regime. Since then the production of foodstuffs 
has diminished, and the population has quadrupled from 9 to 36 million 
(1911). 

At the beginning of the nineteenth century, they feared in Germany, as 
as much as in England, an increase of population. Measures proper for 
restraining marriages were even passed in the legislatures of certain states 
such as Bavaria and Wurtemberg. In order to have the right to marry, 
one had to show sufficient means. Thanks to these restrictive measures 
and to propaganda, the increase of population has remained very slow— 
slower than in France, during a great part of the nineteenth century. Thus 
during the period of thirty-five years, 1847-1871, the number of inhabitants 
increased 13 per cent in Bavaria, 9 per cent in Wurtemberg, while they 
increased 17 per cent in France. 

Events happened which transformed the state of mind, and without doubt 
the faith in the future, without modifying the natural conditions of produc¬ 
tion, and the view changes. During a second period of thirty-five years, 


256 


LUCIEN MARCH 


from 1871 to 1915, the number of inhabitants increased 34 per cent in 
Bavaria, 27 per cent in Wurtemberg, while the proportional increase fell 
to 9 per cent in France. 

A good element of appreciation of the activity and the power of expan¬ 
sion of a people is furnished by the development of its exports, or, if one 
considers 10 states for which one can give at the same time the proportional 
increase of the number of inhabitants from 1875 to 1913 and the relative 
progress of exports, one proves a close relation between the two movements. 


Proportional increase between 1875 and 1913 



POPULATION 

EXPORTS 

France. 

Per cent 

10 

per cent 

80 

Italy. 

29 

145 

United Kingdom. 

45 

160 

Belgium. 

54 

237 

Russia. 

65 

260 

Austria-Hungary. 

38 

383 

German Empire. 

58 

380 

Canada. 

103 

423 

United States. 

138 

386 

Argentine Republic. 

330 

828 



The two series of numbers vary in the same direction. 

IV 

What is to be concluded from these results? Simply that the phe¬ 
nomenon is too complex to be analyzed in its entirety without tracing back 
to elemental facts. 

Let us turn to the father of the family, for it is in fact upon the fathers 
of families that the birth rate of the country depends. We have said that 
this decision depended most generally on three factors: 

1. The expense represented by bringing up a child to the time when 
it can care for itself. 

2. The chances this child has of living effectively, at least in the conditions 
under which its parents have lived. 

3. The opinion of the parents of this expense and these chances. 

Other factors intervene also; considerations of health, well-being, etc., 
but we will only concern ourselves with those which are most general and 
least synthetic. 



















THE WAR AND THE BIRTH RATE IN FRANCE 


257 


Do not regret that, in this grave question, reason is substituted for the 
most simple instincts. Let us force ourselves to see only that which 
commands the true meaning of things. 

At the origin of the problem of the birth rate, one finds two facts of 
economic order and one psychological fact. This last dominates the two 
others, particularly the second. Add that the psychological fact only 
intervenes where the customs and legislation are directed by the sentiment 
of respect for life. For among the primitive people, abortion and in¬ 
fanticide excuse the parents from thinking of the future. They let the 
sexual instinct act freely, for they may cause to disappear the results of 
this action, sometimes, as in Sparta, with the illusory forethought of 
selecting the survivors. 

In our modern society, these procedures are no longer admitted; they 
are supplanted by the prevention of births; that is left to the will of the 
parents who bear the burdens. But this will is guided by judgment and 
sentiment. If judgment is clear and sound, if sentiment is right, the 
voluntary action will be well directed. In the contrary case, it will come 
to evil. But the first condition, in order that the parents be not hindered 
by a too fearful foresight, so that they may act in a sense best conforming 
to the good of society of which they are a part, is that they have a certain 
moral force, that they know how to sacrifice a little of their personal 
interest to the common interest, for maternity always brings some 
sacrifice, at least physical, and that they have confidence in the future. 
One may say that the question of population is above all a moral ques¬ 
tion. A certain optimism is necessary but this optimism ought to follow 
from facts. 

It is always imprudent to ask too much of the sentiment of duty when 
one addresses a whole population. During the war, when invasion roused 
patriotism, it was necessary to impose military service by force. 

So when it is a question of the birth rate. When general education, when 
the comparison of military or economic power of the country shows to all 
families the common duty, nothing better. However, although in this 
matter, no sanction will be legitimate or efficacious, still it will be proper 
to facilitate the accomplishment of this duty. 

That which concerns the care of the future is one of the legitimate pre¬ 
occupations of the head of the family. The movement of general pros¬ 
perity must be such as to make the establishment of children appear easy. 

It is sometimes said that there are fewer children in well-to-do families 
than in poor families. This is true in the sense that if the income of poor 
families increases, the number of their children tends to diminish. But this 
is not really exact for all categories of rich or poor families. 


258 


LUCIEN MARCH 


Let us consider for instance the French statistics of 1906 where the families 
have been classified according to the number of children born in these 
families, whether living or dead; in the families where the marriage has 
lasted twenty-five years or more, the number of children per 100 families is 
equal to 303 among clerks and increases to 360 among their employers, 409 
among laborers, more than 480 among fishermen and sailors of commerce. 

If one classifies the employers of more than 25 years of marriage and aged 
from sixty to seventy years, one finds that the mean number of children 
born in 100 families is only that of 305 in the liberal professions, that it 
increases to 347 in commerce, 370 in agiculture, 385 in the total of industries, 
properly so-called. 

The relative situation of employers in agriculture and industry is not the 
same when one considers the marriages having lasfed less than twenty-five 
years; for the marriages having lasted less than five years, from five to four¬ 
teen years, or from fifteen to twenty-five years, productivity of the mar¬ 
riages is greater in agriculture than in industry. All happens as if the chiefs 
of agricultural exploitations, after having had a determined number of chil¬ 
dren more rapidly than the chiefs of industrial enterprise, stop sooner than 
the latter. 

The detail of professions permits even a distinction between the groups 
of similar industries. The number of children for 100 married men exceeds 
390 in mines and quarries, in the mining and in the textile industries, in 
the enterprises of building and of transportation, while it falls to 350 and 
below in industries of food production, book-publishing, (book-binding or 
printing), goldsmithing, and jewelry. Thus it appears that in the great 
industries the employers have more children and in the small ones fewer. 

Among the commercial professions, the number of children per 100 
families is greatest among the butchers; it is smallest among bankers 
and chiefs of financial enterprise who form a sort of transition between 
industrial or commercial professions and the liberal professions. 

Thus, among employers, productivity seems bound, in a certain measure, 
to the professional characteristics, but these are rather complex: on one 
hand, the intellectuality of the profession, if one may so call it, a small 
productivity, as the number of children per family is small in the liberal 
professions, in the studious professions, in financial enterprises, while the 
manual professions have a productivity relatively higher; on the other hand, 
the chiefs of great industries seem to have a productivity higher than that 
of the small industries and merchants. 

Two factors act in a quasi-independent way; on one hand the intellectual 
character of the professions which leads to late marriages and creates an 


THE WAR AND THE BIRTH RATE IN FRANCE 


259 


environment little favorable to fecundity for reasons which it is not neces¬ 
sary to develop here; on the other hand, preoccupation with the fate reserved 
for the children. In great industries, these will easily find employment for 
their abilities and will obtain without too many difficulties situations equiva¬ 
lent to those of their parents, either in or out of the country. In the little 
enterprises, on the contrary, and except in special instances, such as that 
of butcher, where the employment of the entire family is almost a condition 
of success, the father of the family does not look ahead, without uneasiness, 
to the future laid out for his children. 

Certain of these characteristics will be found among the salaried clerks 
and the laborers. 

The smallest number of children is observed among the clerks of stores, 
waiters in cafes, hotels *and restaurants, office employes and employes of 
public service. 

Among the laborers, the greatest productivity, at least more than 5 
children born in a family founded more than twenty-five years, is among 
smaller laborers and workers in spinning mills. 

One notes that the lace weavers, among which are a great number of 
weavers working at home, have a smaller productivity than the spinners 
(489 per 100 families against 540 among the spinners). One notes also 
that, in agriculture, the domestic workers of the farm, generally lodged at 
the farm, have 395 children per 100 families, while the field workers proper 
have 426. 

But the industries in which the workers have less than 4 children per 
family are numerous. Those who have about 350 children per 100 families 
founded more than 25 years are makers of wooden shoes, coopers, toymakers, 
saddlers, tailors, printers, metalworkers, electricians, jewelers, and silver¬ 
smiths, various workers in commerce, drivers, deliverymen ( livreur”). It 
seems that professions of smaller industries, and especially professions 
in cities,, give the smallest figures. For the masons, day laborers, people 
without profession, generally occupied in the cities, one counts 464 children 
born per 100 families; among the workers of industrial service of the State, 
roadmenders, etc., the productivity exceeds 390 children born per 100 
families; it decreases to 360 among the police, and customs employes, etc., 
to 350 for workers and under agents of the post and telegraph. 

Finally, among personal servants, it decreases to less than 3 children 
born per family, always for the heads of families married more than twenty- 
five years. 

On the whole, among laborers, and workers in great industries where the 
work is relatively regular and abundant, when the agricultural work offers 


260 


LUCIEN MARCH 


a real stability, when the dwelling is either in the country, or among industrial 
collections grouping the laborers of the same class, productivity is relatively 
high; it is lowest where the small artisans live most, in the trades carried on 
in cities, also where profession demands to the least degree physical force. 
It is also small where the persons classified as workers are confined to the 
category of clerks and especially where the conditions of employment, the 
conditions of lodging, make preferable households without children or with 
few, rather than households burdened with children. 

From the total of the preceding statements we will remember that if 
the workers in general have more children than the employers there are 
not lacking professions where they are fewer. In the second place, for one 
as for the other, it is the great industries which seem more favorable to 
productivity, and small ones less favorable. Naturally here the influence 
of environment exercises a certain influence, the regions of great industry 
are generally other than those of small industry. 

The preceding observations (the pictures shown in the exposition rooms 
of the Congress illustrate these observations) confirm, although not entirely, 
those that have often been made on the relation between fertility and social 
standing. This being at once the function of income and education, those 
are the most fortunate categories where education is the most refined, or 
where the number of children is the most limited. On the contrary, fer¬ 
tility would be greatest in the poorest environments, in those where the 
kind of life is the plainest. 

If, in a general way, there is in this observation a great part of truth— 
the comparison of districts of great cities according to categories classified 
according to exterior signs of income show it—there are modalities which 
must be taken into account. No doubt, for example, that employers are 
generally more fortunate than their employes, and meanwhile they have 
notably more children than the latter. On the other hand, employes, 
who receive generally higher wages than laborers, have fewer children than 
the latter. 

The question has often been studied, and it is important that new con¬ 
tributions be brought to it. 

We will borrow for new indications recent statistics of France drawn up 
by the aid of family schedules filled out in 1907 by a great number of em¬ 
ployes and workers remunerated by the budgets of the State, departments 
and communes (Conseil superieur de statistique, bulletins 10 et 11 Statis- 
tique Generale de la France, Statistique des families en 1906). 

These functionaries have been classified according to the annual showing 
of the actual emoluments and, considering only those whose marriages have 


THE WAR AND THE BIRTH RATE IN FRANCE 


261 


lasted more than 15 years, the number of children born per 100 families 
has been calculated: 


ANNUAL SALARY IN FRANCS 



500 at 
most 

501 to 
1,000 

1.001 

to 

1,500 

1,501 

to 

2,500 

2,501 

to 

4,000 

4,001 

to 

6,000 

6,001 

to 

10,000 

More 

than 

10,000 

Aver¬ 

age 

Marriages lasting fifteen to twenty-five years 

Clerks. 

277 

241 

259 

245 

223 

231 

239 

238 

237 

Laborers. 

329 

321 

293 

280 

254 

234 



307 

Marriages lasting more than twenty-five years 

Clerks. 

330 

301 

305 

280 

264 

264 

261 

286 

285 

Laborers. 

348 

363 

346 

329 

305 

240 



385 


All classes taken together, the numbers which precede are in accord with 
those which have been determined with the aid of the general census, 
either for clerks or for laborers or sub-agents of the public service. 

Comparing now the numbers of children by classes of salaries, one will 
note that, among the laborers, the number of children diminishes regularly 
as the salary increases; among the clerks it diminishes until it reaches a 
minimum for clerks earning 2500 to 10,000 francs per year; it rises for 
clerks whose annual payment exceeds 10,000 francs. 

To complete these proofs, it is proper to remark that salaries and emolu¬ 
ments depend in great measure on the region or settlement where each 
clerk or laborer lives. Change in fertility is submitted to a double influence, 
showing that salary only partly determined. 

The influence of environment has been evident in observing in all France 
the families of limited classes of agents scattered in all the territory, 
generally in the rural communes, the roadmenders and the gardes-champ&tre 
(rural police?). 

For these agents, fertility is analogous to that of the population in the 
midst of which they live, greater in the regions of high birth rate, smaller 
in the regions with a low rate. 

One has preceded with a similar research among the clerks properly so 
called of prefectures and mairies. The personnel of the employes (not 
composed of boys, guardians, laborers, etc.) has, in general, fewer children 
proportionately as the number of inhabitants of the city has increased. 
It is the same for the populations of these cities. But, if one establishes the 
relation between the fertility of these functionaries and general fertility, 
one remarks that the first is less variable than the second. 








































262 


LUCIEN MARCH 


In 1901, 100 families founded more than 15 years had 199 surviving chil¬ 
dren in Paris, 228 in cities of more than 500,000 inhabitants, 266 in the 
smaller cities. Among the administrative employes, the corresponding 
numbers are 183,198,215, or 92 per cent, 87 per cent, 81 per cent of the pre¬ 
ceding. Employes have in some degree a specific fertility which depends 
less on environment than that of laborers. 

One obtains results analogous to the preceding, when one determines the 
proportional number of sterile families. 

Among the marriages having lasted more than 25 years, the number of 
sterile marriages to 1000 marriages varies as follows, showing the annual 
income: 


ANNUAL SALARY IN FRANCS 



Less 

than 

1,000 

1,001 

to 

1,500 

1,501 

to 

2,500 

2,501 

to 

4,000 

4,001 

to 

6,000 

6,001 

to 

10,000 

More 

than 

10,000 

Clerks. 

95 

86 

99 

113 

101 

111 

109 

Laborers. 

70 

74 

91 

98 

100 





Proportional number of families having had more than 7 children 


Clerks. 

56 

53 

41 

33 

26 

23 

52 

Laborers. 

95 

86 

75 

55 

50 





On the whole, the statistics of French families permit one to see in what 
measure fertility is bound up with the social situation. Numerous factors 
intervene: for instance, the chiefs of enterprises in the most industrial 
regions of the country: the North, the region about Lyons—have many 
children, more children per family than many other less fortunate classes. 
Among the laborers, the miners of Pas-de-Calais have likewise many 
children in relation to other laborers. In these two cases the parents have 
no fear as to the future of their children. The great employer knows that 
he can easily establish his; the mine laborer knows that there will always 
be work in the mine for his. 

This sentiment becomes general when one perceives continued progress 
everywhere, in the agricultural, industrial and commercial movements, and 
in the action of public powers in favor of education, apprenticeship, exporta¬ 
tion, emigration, great works. 

Confidence in the future is then assured. It is this factor which seems 
to have played an important role in Germany after the constitution of the 
Empire and the war of 1870. But the two factors which we have just 
considered, a certain courage on the one hand and a certain optimism on 



































THE WAR AND THE BIRTH RATE IN FRANCE 


263 


the other, do not suffice always; it seems useful to ward off at first the ob¬ 
stacles which we have recognized, that is to say, to lighten the burdens 
which the maintenance of children cause parents. 

Here it is proper to proceed with method. Since it is a question of finan¬ 
cial participation, it is expedient to place the effort where it is most necessary 
and to seek to obtain the maximum result from the sums used. 

It is humane to seek that the children brought into the world be raised 
under the best conditions for health. 

It is good not to go against the natural course of things to limit oneself 
to bringing simply the spark which puts the fire to the pile. 

v 

These considerations tend to favor the birth rate in the country. It is 
there that depopulation is raging, not that the birth rate is less elevated than 
in the cities—the contrary is true—but because of the emigration from the 
country to the city. 

One notices this when one compares the movement of the number of 
inhabitants in the French censuses of different periods, either in urban or 
rural population. 

In 1856 the rural population was 26 million of the 36 million inhabitants 
in all France; in 1911, the number has fallen to 22 million, while the total 
population had increased to almost 40. Also the urban has been consider¬ 
ably augmented, passing from 9,800,000 inhabitants in 1856 to 17,500,000 
in 1911; it almost doubled. It has doubled also in the class of cities of more 
than 10,000 inhabitants. 

It is useless consequently to seek to increase the population of cities by 
artificial means since they increase so rapidly alone, until there is a veritable 
overcrowding in great cities. But it is necessary to increase that of the 
country for reasons of hygiene, social stability, good economy also, for it is 
there that children cost least. 

It is there also that the birth rate is already the highest, where one will 
find families best disposed to have numerous children. One states that the 
birth rate increases in proportion to the altitude. But, in France at least, 
it is from the high altitudes that have come the strongest current of 
emigration. 

Children cost much less to raise in the country than in the city; ifiis in 
the country that poverty is the most disquieting, which ought to interest 
the farmers to assure themselves of the number of children capable of aiding 
them by their work; it is there that the growth of children takes place under 
the best conditions of health, especially if a system of maternal education 


264 


LUCIEN MARCH 


is instituted; it is there that one is near the foundation of the population, 
and where marriages are made with full knowledge of antecedents. Even 
as one rejuvenates trees from the stump, so the renewing of the population, 
necessary to combat retrogression, ought to be worked from the base. The 
best always come from a vigorous stock, as the best fruits and the most 
beautiful flowers spring from well grafted roots. In the cities, national 
effort ought to tend to improve transportation to facilitate rapid communi¬ 
cation which will permit the largest extensions outside the crowded areas. 

In France a law of July 14, 1913, gives to every family which has at least 
three children less than thirteen years a monthly allotment for each child 
beyond the third under thirteen years, while the child is living and has not 
reached the age of thirteen years. The communes, the departments and 
the State share the expense. 

Another law, that of June 28,1918, gives an important share of the state 
(power) to the departments which allow encouragements to birth. This 
participation varies in inverse ratio to the richness of the department and 
in direct ratio to the number of families having more than four children. 
It carries at the same time the useful premium for the maintenance of 
children and the premium of foresight destined to assure a life-annuity to 
old parents or a capital to grown children. 

In cities and industrial centers, numerous patronal associations have been 
formed to assure to laborers and clerks allotments varying according to the 
number of children. The treasury is kept filled by payments of chiefs of 
enterprise proportional to the salaries paid by each one of them. Thus the 
industrial has no interest in employing a bachelor any more than the head 
of a family. 

The employes of the State and those of great private enterprises receive 
the same family allotments added to their salaries. 

Finally, the fiscal legislation assures important exemptions to heads of 
large families and a surcharge to bachelors and families without children. 

The tariffs of income tax—impots cedulaires et imp6ts globa—take 
account of the number of children; impot globa surcharges the bachelors as 
well as married men without children. The inheritance taxes grant reduc¬ 
tions according to the number of children living or represented, and sur¬ 
charges when the defunct has left no children. Reductions are given on the 
railroads to members of families which have many children. 

A severe law has been promulgated, July 31, 1920, against abortion and 
the sale of contraceptive measures. 

An important movement thus exists in France which cannot but be favor¬ 
able to increasing the birth rate. None of the measures adopted offer 
dispositions contrary to the legitimate exigencies of eugenics. 


THE WAR AND THE BIRTH RATE IN FRANCE 


265 


Let us add that the struggle against tuberculosis and the effects of venereal 
disease have gained much activity since the war; numerous dispensaries 
have been erected, to such a point, that in spite of the increase of these 
diseases, one cannot find, as might have been feared, an increase of the 
special disability of children, excepting naturally those who were born or 
who passed their childhood in the regions invaded by the enemy. 

The decline of the birth rate is a phenomenon which has shown itself in a 
great number of countries. The intensity of the movement is very different 
in different States; its effects depend, in great part, on the long or short dura¬ 
tion of time since the phenomenon commenced to appear. The causes 
are almost the same everywhere; the means of combating the causes are 
not known to be very different, although the action of moral influences 
depends naturally much on general mentality. As to the other influences, 
the experience which has been had in France cannot fail to be instructive 
for all nations and for all those who are interested in this still conjectural 
science known as Eugenics. 


THE EFFECTS OF INBREEDING ON GUINEA-PIGS 
SEWALL WRIGHT 

Bureau of Animal Industry, United States Department of Agriculture, Washington, D. C. 

An experiment on the effect of inbreeding, using guinea-pigs as material, 
has been carried on since 1906 by the Bureau of Animal Industry of the 
United States Department of Agriculture. 

Twenty-three families of guinea-pigs were maintained for a number 
of years by mating exclusively brother with sister among the descendants 
of twenty-three original pairs. In sixteen cases both of the original 
parents came from a stock which had already been maintained for twelve 
years without the infusion of fresh blood, by the Experiment Station of the 
Bureau of Animal Industry. In the remaining families the original females 
came from the above mentioned stock, while the original males were pur¬ 
chased from a local dealer. 

In 1911 a number of animals were selected from the stock of the Experi¬ 
ment Station to start a control experiment. The mating of even second 
cousins has been avoided in this stock, which has been called Experiment B. 

Eighteen of the inbred families were still on hand in 1916, having then on 
the average about eleven generations of brother-sister matings back of them. 
At this time most of the families were disposed of in order to make room for 
cross-breeding experiments and to obtain larger numbers from the five 
families which it was decided to retain. These are families, 2, 13, 32, 35 
and 39. They were retained in part because they already occupied many 
pens, but largely because of the possession of contrasting characteristics in 
size, fertility and coat pattern. Each of these families is at present very 
homogeneous in heredity. Family 2 is descended wholly from one mating 
in the sixth generation of inbreeding. Families 13, 32, 35 and 39 come from 
single matings in the seventh, eleventh, twelfth and eighth generations 
respectively. 

The total number of animals involved in the experiment on inbreeding 
and cross-breeding has been about 35,000. 

THE EFFECTS OF INBREEDING ON VIGOR 

It is noteworthy that there has been no very obvious decline in vigor, 
although the families are now on the average in the sixteenth generation 

266 


EFFECTS OF INBREEDING ON GUINEA-PIGS 


267 


of brother-sister mating and one of the most vigorous (no. 35) has reached 
the twenty-third generation. 

There has, however, been some decline in vigor in all respects which 
have been studied. The decline is most marked in fertility, including 
both frequency and size of litters. It has been so great in this respect 
that it would have to be recognized even though the decline in other 
respects were assumed to be due wholly to less favorable environmental 
conditions. 

That there has been a real genetic decline in the inbred stock in all ele¬ 
ments of vigor is shown by comparison with the control stock B, which has 
been superior in every respect. Still better evidence has been obtained by 
comparison of the inbreds with the young from crosses between the different 
families raised at the same time and under the same conditions. 

THE EFFECT OF CROSSING 

In interpreting the effects of crossing, the characteristics which depend on 
the hereditary make-up of the young must be distinguished from those which 
depend on the dam or sire. In studying these questions, inbred females 
have been mated with inbred males of another family (experiment CO) 
and with crossbred males (experiment CA). Crossbred females have been 
mated with brothers (experiment Cl), unrelated crossbred males (experi¬ 
ment CC) and inbred males of an unrelated family (experiment AC). 

Size of litter appears to depend wholly on the dam. There is little or no 
improvement in the experiments in which the dam is inbred (CO, CA). 
There is, however, a marked increase, 28 per cent, in the litters produced 
by crossbred females of experiment AC. 

In experiments Cl and CC in which the dam was also crossbred, there was 
an increase, but a smaller one (10 to 14 per cent). The influence on size of 
litter, of the greater frequency of litters in these experiments, is believed to 
be responsible for the smaller increase then in experiment AC. 

The frequency of litters depends to a greater extent on the sire than on the 
dam. This is shown by the greater increase in frequency over the inbred 
stock when a crossbred male is mated with an inbred female (CA) as com¬ 
pared with the mating of an inbred male with crossbred female (AC). Ex¬ 
periment CA produced 19 per cent more litters per year per mating than the 
inbreds, while AC produced only 9 per cent more. When both parents are 
crossbred there is a still greater improvement (36.1 and 33.3 per cent in 
experiment CC and Cl respectively). 

The percentage of the young born alive depends almost wholly on the 
dam. There is little or no improvement in experiments CO and CA, but 


268 


SEWALL WRIGHT 


an increase of 6 to 8 per cent, where the dam is crossbred. The percentage 
which are raised to 33 days of age of the young born alive depends both on 
the dam and on the heredity of the young. There is a marked increase, 9 
to 12 per cent, in all of the crossbreeding experiments mentioned above. 

Somewhat similarly, birthweight depends largely on the dam, while 
the gains between birth and thirty-three days depend to a considerable 
extent, though far from wholly, on the heredity of the young. Guinea pigs 
become independent of the dam at a very early age. There is an increase of 
2 or 3 per cent in experiments CO and CA, but one of about 7 to 10 per cent 
where the dam as well as the young are crossbred. In the gain between 
birth and thirty-three days, there is an improvement of about 13 per cent 
in the first cross, which is somewhat increased in the young produced by 
crossbred dams. There is an increase of 12 per cent in the adult weight in 
the first cross (CO). This increase is lost where the parents, though cross¬ 
bred, are brother and sister (Cl). The influence of the dam does not appear 
to extend to the adult weight. 

A loss in the improvement brought about by the crossing becomes ap¬ 
parent in the second generation of inbreeding following a cross (experiment 
C2) in those cases in which it is not apparent in the first generation. 

COMPARISON OP DIFFERENT FAMILIES 

A comparison of the different inbred families with each other has revealed 
persistent differences in color, pattern, tendency toward polydactylism, 
tendency toward production of monsters, mortality among the young, 
weight and both elements of fertility. It was found that the differences in 
these respects could not be interpreted merely as differences in general 
vigor. Vigor above the average in one respect was as likely as not to be 
found associated with a subnormal record in another respect, the correla¬ 
tion between the records of the families in two respects coming out in most 
cases substantially zero. 

EXPLANATION OF THE RESULTS OF INBREEDING AND CROSSING 

These results harmonize well, on the whole, with those found by other 
investigators. It is believed that they can be explained as consequences 
of the current theory of heredity without recourse to the rather mystical 
ideals which once prevailed in regard to inbreeding. There appear to be 
independently inherited factors which affect frequency and size of litter, 
ability to bear the young successfully, vitality and growth as well as for 
color, pattern and the other characters in which the families differ. There 


EFFECTS OF INBREEDING ON GUINEA-PIGS 


269 


seem to be surprisingly few factors which act on all of these characteristics. 
The concept, hereditary vigor, thus becomes merely an expression for the 
sum of a number of independently inherited qualities and not an entity. 

The factors which cause reduced vigor in any respect appear to be in 
general recessive. The primary effect of inbreeding is to render homozygous 
a random group of the factors present in the original stock. Some combina¬ 
tion of factors, good, bad and indifferent, thus becomes fixed in each inbred 
line. As the recessive factors, tending toward lack of vigor, are as likely to 
become fixed as the dominant ones there is on the average a decline in 
vigor in each respect. Moreover, owing to the likelihood that many fac¬ 
tors for vigor will be linked genetically with factors causing weakness, it is 
to be expected that vigor in all respects will be found in very few lines, even 
where there is careful selection. In the present case, there was very little 
conscious selection but a considerable amount of natural selection was, of 
course, unavoidable. 

On crossing two inbred lines, each, as a rule, supplies some of the domi¬ 
nant factors lacking in the other, with the consequence that there is increased 
vigor in so far as the character in question depends on the heredity of the 
animal itself. In the next generation, if brother-sister matings are made, 
there should be a decline as compared with the first generation in characters 
which depend wholly on the animal itself. The decline from this cause may, 
however, be balanced or more than balanced by the improvement due to the 
influence of the crossbred-dam. 


IS INBREEDING INJURIOUS? 


HELEN DEAN KING 
The Wistar Institute of Anatomy and Biology 

There is a widespread belief that close inbreeding is necessarily injuri¬ 
ous, and that it leads to a loss of body size, of constitutional vigor and of 
fertility, accompanied by a greater susceptibility to disease and by a tend¬ 
ency to malformations. 

If we seek in the older literature for the basis of the prejudice against 
inbreeding that has existed for many centuries we find much that is myth 
and superstition, very little that has scientific value today. The first 
serious attempt to study this problem was made by Charles Darwin (78) 
who inbred various species of plants during a period of eleven years. Dar¬ 
win’s work aroused much interest among zoologists and led to several series 
of inbreeding experiments with mammals, of which the most important were 
those of Crampe (’83, ’84), and of Retzima-Bos (’94) with the rat. The 
results of all of this work seemed to justify the prejudice against inbreeding, 
and for thirty years have been the classic examples of the harmful effects of 
this form of mating. 

With the recent development of the science of Genetics there has come a 
renewed interest in the problem of inbreeding, and during the past few years 
a number of important investigations have been carried on which show that 
there is no physiological law forbidding inbreeding and that the results 
obtained, whether good or bad, can readily be interpreted according to 
Mendelian laws. 

For some time I have been conducting a series of inbreeding experiments 
with the albino rat, and I desire to give a brief account of this work in order 
to show the effects of close inbreeding in this animal when the experiments 
are carried on under controlled conditions for a relatively long period of 
time. 

The experiments were begun in 1909 with a litter of four albino rats, two 
males and two females, taken from a stock colony maintained for research 
purposes. From these two pairs of rats two lines of inbred animals have 
descended that have been kept under the same conditions of environment 
and of nutrition and continued for forty generations through the mating of 
brother and sister from the same litter, this being the closest form of in- 

270 


IS INBREEDING INJURIOUS 


271 


breeding that is possible in mammals. At the present time the record for 
this series comprises some 7000 litters containing over 50,000 rats. 

There has been a very rigid selection of animals to be used for breeding 
purposes throughout the course of this investigation. Small, weak, de¬ 
fective individuals have been eliminated, either at birth or before reaching 
sexual maturity, and only the largest and most vigorous animals in the large 
number available have been used to continue the inbred lines. 

The first six generations of these animals showed all of the characteristics 
that are popularly supposed to result from close inbreeding. There was a 
steady decline in the vigor of the animals in succeeding generations; many 
females were sterile, and those that did breed produced small litters that 
contained many stillborn young; most of the animals were undersized; 
and a number showed malformations, particularly deformed teeth. These 
results agreed with those obtained by Crampe and by Retzima-Bos, and 
seemed to indicate that in the rat close inbreeding soon leads to marked 
physical degeneration (King, 18). 

Fortunately for this work it was discovered that the stock albinos in the 
colony that were not inbred were exhibiting the same evidence of degenera¬ 
tion as the rats in the inbred strain. It was obvious, therefore, that the 
unfavorable condition of the animals could not be due to inbreeding alone. 
We found that the rats were suffering from malnutrition due to the character 
of the food they received. When the food was changed there was at once 
a very decided improvement in the condition of the rats: they gained in 
size and in weight; there was a marked increase in fertility and in general 
vigor; malformations soon disappeared, and not to exceed a dozen abnormal 
individuals have been found among the many thousands of inbred animals 
that have been born since that time. 

Since inbreeding is believed to have its most marked effects on body size, 
on fertility and on constitutional vigor, these are the characters that will be 
considered mainly in summarizing the results of these experiments. 

In each generation after the seventh the individuals in from three to five 
litters of each of the inbred lines were weighed at stated intervals from birth 
until they were fifteen months of age. As controls for the inbreds, series 
of outbred stock Albinos were reared at various times under the same con¬ 
ditions of nutrition and of environment as the inbred rats, and they were 
weighed at the same age periods. 

Figure 1 shows graphs for the growth in body weight of males belonging in 
the seventh to the fifteenth generations of the inbred strain, and for males in 
the series of outbred stock controls. 


272 


HELEN DEAN KING 


During the first month of postnatal life there was little difference in the 
rate of growth of the inbred and of the stock animals, but after this time, as 
figure 1 shows, inbred males increased in body weight much more rapidly 
than the stock males. When the animals were at their prime, at about 
eight months of age, the inbred males were about twelve per cent heavier 
than the control males. 



Fig. 1. Graphs showing the weight of the body with age for 156 males belonging in 
the seventh to the fifteenth generations of the inbred strain, and for 50 males in a series 
of stock controls (Data: King,’18). 

The female albino rat is normally a much smaller animal than the male of 
like age and is less variable in body weight. Figure 2 shows graphs for the 
body growth of inbred and of stock females that belonged in the same litters 
as the males whose growth graphs are given in figure 1. 

The graphs in figure 2 show that in the adult state females in the seventh 
to the fifteenth generations of the inbred strain were heavier animals than 
the females in the control series, although the difference in weight at any 
age period was not as great as in the case of the males. 



IS INBREEDING INJURIOUS 


273 


The present status of inbred animals as regards their body weight at 
various age periods is shown in figure 3 which gives graphs for the body 
growth of males belonging in the thirty-sixth to the thirty-eighth genera¬ 
tions of the inbred strain and for males in the series of stock controls. 

Inbreeding has not, as yet, decreased the body size of these rats to any 
appreciable extent, since the course of the graph for the body growth of 
males in later generations of the inbred strain runs practically the same as 
that for the males of the earlier generations (cf. figs. 1 and 3). For some 
unknown reason this series of stock controls was inferior to the general run 



Fig. 2. Graphs showing the weight of the body with age for 169 females belonging in 
the seventh to the fifteenth generations of the inbred strain, and for 50 females in a series 
of stock controls (Data: King, ’18). 

of stock Albinos, both in body size and in fertility. The average body 
weights of adult stock males at various age periods were about 20 per cent 
less than that of inbred males of the same age (fig. 3). Graphs for the body 
growth of the females in these two series would show that the inbred 
females were much heavier animals than the controls at every age period. 

The fertility of the animals in the inbred strain can best be shown by 
comparing the average size of inbred litters with that of litters cast by 
out-bred stock females reared under the same environmental conditions as 
the inbred rats. 




274 


HELEN DEAN KING 


In the first twenty-five generations of the inbred strain, as table 1 shows, 
the litters contained an average of 7.39 young, while the litters of the stock 
controls averaged only 6.75 young: In the twenty-sixth to the fortieth 
generation, with the data for the last generation incomplete, the inbred 
litters were as large as those in the earlier generations, but the litters of the 
control series were relatively much smaller. In the inbred strain as a whoje, 
therefore, there has been no marked decrease in fertility as a result of con¬ 



tinued close inbreeding, and the litters have been larger, on the average, 
than those of stock Albinos reared as controls. 

The so-called constitutional vigor of any animals may be tested in a 
variety of ways: for example, by agility and by mental alertness, by longevity 
and by ability to resist disease. In the two latter respects inbred rats 
compare very favorably with stock animals, Psychological tests showed 
that inbreds of the early generations that were suffering from malnutrition 



IS INBREEDING INJURIOUS 


275 


were very inferior to stock animals, reared under more favorable environ¬ 
mental conditions, as regards their ability to form and to retain habits (Bas¬ 
set, ’14), and autopsies showed that these rats had a very low brain weight. 
Inbreds of the twelfth to the fourteenth generations were found to have a 
normal brain weight and to be but slightly inferior to stock animals in their 
behavior reactions (Yerkes, ’16; Utsurikawa, ’17). No tests have been 
made of animals in later generations of the inbred strain, but autopsies of a 
number of individuals of the thirty-eighth generation showed that 75 per 
cent of them had a brain weight heavier than the standard weight for stock 
animals of like age. 

The results of these experiments, taken as a whole, indicate that in the 
rat, inbreeding is not necessarily injurious, even when continued for forty 
generations of brother and sister matings—a period that, assuming three 
generations to a century, would cover 1300 years of human life. 

TABLE 1 


Showing the average size of the litters in series of inbred and of outbred albino rats 


INBRED ALBINO RATS 

OUTBRED STOCK ALBINO RATS 

Generations 

Number 

litters 

Number 
’ individuals 

Average 
number young 
per litter 

Number 

litters 

Number 

individuals 

Average 
number young 
per litter 

1-25 

1898 

14,029 

7.39 

260 

1756 

6.75 

26-40 

2047 

15,084 

7.37 

254 

1583 

6.23 

1-40 

3945 

29,113 

7.38 

514 

3339 

6.49 


Mendelism has shown that, under normal conditions of outbreeding, 
characters which are favorable for the species tend to dominate in heredity 
those which are harmful. The genetic factors for the injurious characters 
persist in the germplasm from generation to generation, however, and bring 
out the latent, undesirable traits whenever conditions permit. Outbreeding, 
therefore, tends to hide defects but it does not eliminate them. 

Inbreeding invariably brings to light the latent characters that were hid¬ 
den by outbreeding: it cannot, from its very nature, introduce any new 
characters into the stock. Since the constitution of individuals produced 
by inbreeding depends upon the chance segregation and recombination of 
genetic factors already existing in the stock when the inbreeding was begun, 
some inbred individuals will exhibit the latent, undesirable traits previously 
hidden by outbreeding: others will show favorable qualities. Random 
matings in such an inbred stock will not suppress the undesirable traits; 
but if only the superior individuals are allowed to breed, the unwanted 




















276 


HELEN DEAN KING 


traits in the stock can gradually be eliminated if linkage does not exist, 
and they will not reappear unless through mutation, since the germplasm 
of the stock will no longer contain the genetic factors upon which these 
traits depend. 

Inbreeding, with selection, thus becomes a powerful agent to purify a 
stock, to bring about a concentration of desirable traits, and to eliminate 
serious defects. It is through inbreeding, combined with skillful selection, 
that the most celebrated breeds of cattle, of sheep, of horses and of dogs 
have been evolved. 

In spite of his great superiority, man is subject to the same laws of 
heredity as the rest of the organic world. Not only feeble-mindedness, 
epilepsy and insanity, but also general mental efficiency and marked ability 
in music, in art and in literature are undoubtedly transmitted from genera¬ 
tion to generation according to the same Mendelian laws that govern the 
inheritance of the color of the hair and of the eyes (Davenport, ’ll). The 
effects of consanguineous marriages on the offspring of man can also be ex¬ 
plained by Mendelian laws as readily as can the effects of inbreeding in 
lower mammals. 

The germplasm of man, as it is constituted at the present time, probably 
contains genetic factors for a large number of latent defects. In the marri¬ 
age of non-kin these defects, as a rule, remain hidden in the offspring, 
since genetic factors for the same defects are not usually present in the germ¬ 
plasm of both parents and therefore the normal character dominates the 
defective. The danger in the marriage of near kin lies in the possibility that 
the germplasm of each parent will contain factors for the same defect. 
Under these conditions the offspring gets a double “dose” of the trait held 
by both parents in common and therefore shows the defect, often in an 
intensified form. If, however, the same congenital defect or undesirable 
trait does not appear in the three previous generations of two cousins, in¬ 
cluding collaterals, there is no more danger that the offspring of such 
marriages will be defective than there is danger of the appearance of de¬ 
fective children in any family (Davenport, ’ll). 

History teaches that the prejudice against consanguineous marriages 
that has persisted from the beginning of the Christian era until the present 
time did not exist among the early nations, and that such marriages were 
common for many centuries among the Greeks, Phoenicians, Jews, Peru¬ 
vians and Egyptians. One of the longest of known human pedigrees, 
that of the royal Ptolemies of Egypt, is noted for its close inbreeding, brother 
and sister marriages being very frequent. There is no evidence that con¬ 
sanguineous marriages were injurious to any of these early nations. The 
decline of the Greeks and of the Egyptians came when they ceased to be 


IS INBREEDING INJURIOUS 


277 


an exclusive people and the strength and vigor of the race were sapped by 
vices and luxuries introduced from other countries. 

In the present status of society there is but little thought for the well 
being of the generations yet to come. Marriages are practically unre¬ 
stricted; known transmittable defects in families are usually ignored: and 
the unfit are freely allowed to reproduce their kind and add to the increasing 
number of defectives that are a burden and a menace to the state. Laws 
forbidding consanguineous marriages are therefore both desirable and neces¬ 
sary, since there are comparatively few families that are known to be free 
from serious hereditary defects. 

Sometime in the future an enlightened people will appreciate the value of 
favorable mutations and of combinations of genetic factors that produce 
unusual ability, and will endeavor to implant them in the race through con¬ 
sanguineous marriages, just as the expert breeder of today has taken ad¬ 
vantage of the appearance of desirable traits and fixed them in his stock 
through the skillful use of inbreeding and selection. When the time comes 
that marriage is based not only on the physical fitness of the individuals but 
also on their recorded pedigree for several generations, and is absolutely 
forbidden to the unfit, the surest way of improving the race will be through 
consanguineous marriages in families in which the members show excep¬ 
tional mental and physical endowment in ways that are of value to them¬ 
selves and to the community at large. Many of the ills to which man is at 
present subject, for example epilepsy, will then vanish; superior and desira¬ 
ble traits will appear in an ever increasing number of individuals and in 
time become the heritage of the race. 

REFERENCES 

Basset, G. C. 1914 Habit formation in a strain of albino rats of less than normal 
brain weight. Behavior Monographs, vol. 2. 

Crampe, H. 1883 Zucht-Versuche mit zahmen Wanderratten I. Landwirthschaft- 
liche Jahrb., Bd. 12. 

Crampe, H. 1884 Zucht-Versuche mit zahmen Wanderratten II. Landwirth- 
schaftliche Jahrb. Bd. 13. 

Darwin, Charles 1878 The Effects of Cross and Self Fertilization in the Vegetable 
Kingdom. Second edition, London. 

Davenport, C. B. 1911 Heredity in Relation to Eugenics. New York. 

King, Helen Dean 1918 The effects of inbreeding on the growth and variability in 
the body weight of the albino rat. Jour. Exper. Zool., vol. 26. 

Ritzema-Bos, J. 1894 Untersuchungen uebef die Folge der Zucht in engster Blutver- 
wandtschaft. Biol. Centralbl., Bd. 14. 

Utsurikawa, N. 1917 Tempermental differences between outbred and inbred strains 
of albino rats. Jour. Animal Behavior, vol. 7. 

Yerkes, A. W. 1916 Comparison of the behavior of stock and inbred albino rats. 
Jour. Animal Behavior, vol. 8. 


INTERMARRIAGE OF BLOOD RELATIVES IN THREE OLD NEW 
ENGLAND COMMUNITIES 

RUTH MOXCEY MARTIN 
Woodbury , Connecticut 

The town of X-is one early colonized from England. Some eight 

or ten families were the original settlers. They were of the typical sturdy 
pioneer sort, with the love of home, a pride in family and during the Revo¬ 
lution a marked tendency to be British sympathizers (though not all were 
out-spoken Tories). 

X-is a coast town so situated that both fishing and wooden ship 

building are practicable. These pioneers were compelled to be their own 
tradesmen, their own agriculturists, etc., in true pioneer fashion. Today 
one finds the fine old names in all trades and occupations in the town. It 
matters not whether you be a barber or the editor of the town paper, the 
boarding-house keeper or acknowledged hostess with weekly “at home” 
day—if you bear one of the original settler’s surnames you belong. The 
boarding-house mistress, if her name and blood be of the elect, may serve 
boiled dinner to any number of day laborers, rank outsiders though they be, 
and yet later be of those who sound the old iron knocker to have afternoon 
tea with the judge’s wife at the big house at the end of the street. Today, 
as before the Revolution, the bulk of the surnames of the town are confined 
to the eight or ten original when the grant was settled. This shows what 
records and townspeople affirm to be true that intermarriage has been 
marked. 

There are today many inhabitants with incomes, many intelligent and 
delightful people. The town can show many graduates of well-known 
universities, colleges and metropolitan conservatories of music. There is a 
noticeable heredity of musical skill frequently linked with erratic tempera¬ 
ment. There are writers of verse and prose; various men of unusual interest 
in some of the finer arts of civilization. Instances are: one townsman with 
a marked passion for modeling in clay and another had achieved noticeable 
success as a naturalist. But in this town so proud of ancestry, so certain 
that blood and only blood counts, there lurks a certain dread, a fear which 
you can call out at will in the faces of its townsmen if you talk with them 
of their genealogy. Possibly a conversation with an old sea captain will 

278 




INTERMARRIAGE OF BLOOD RELATIVES 


279 


give as vivid a picture of this as any lengthy set of statistics. He is a man 
noted in his town for quiet, pleasant ways, even normal in his life and 
reasonable success. 

“ So you wish to hear all about my family from the first of the name who 
settled here and you wish to know what traits of ours have come down 
through the generations!” Then, with a bitter curl of his fine lips (he 
knew the town clerk's records had been well canvassed) “I might just as 
well be frank at the first. I suppose the principal thing this house over 
your head has seen (and it has been the roof tree of six generations of my 
ancestors) is suicides. Nor is this house the only one hereabout of which 
it would be equally true." Then with a keen look at the field worker, 
“Had you heard this is called suicide town? Is that what brought you 
here? Well, for towns about it is how we are known. My grandmother 
hung herself in the attic here; her father, his sister, one of my own uncles, 
two aunts on the other side of the house, and one of my brothers all left 
this world because they willed to. You can find it set down over there in 
the town records. Yes, yes, it is suicide town!"—and his whole bearing 
showed the oppressing weight of such facts to his finely sensitive and doubt¬ 
less somewhat morbid nature. For a moment we were silent. Then once 
more in answer to the question as to what in his opinion caused this, he 
began to speak. “What did it? How came it? Pride, miserable pride, 
that’s what and I’ve been brought up to it and I feel it and most of the time 
I just take it for granted. But I’ve thought and thought. Years and years 
ago it was just the same. I’ve heard my great grandfather say it and 
other old people and folk in the family on down, that no families along 
this coast were anywhere near of so fine blood as we. (I mean in any of 
the little towns.) They believed it and lived it only to their own and our 
undoing. ‘Remember blood tells’ and from children we were made to 

know that no one was good enough to marry who wasn’t a B-or an 

L-or-(and he proceeded with the list of surnames before re¬ 

ferred to). I’ve heard that years ago when young folks ever did marry 
outside it was made so miserable for them that they left the town in time. 
First of all it was to keep property here. They had something when they 
came here, our sires. There is old table silver and such here in town yet to 
show they were real fine folk. Then after they’d done so a few generations 
it was the thing to do. We here did marry in with one town just south 
but some of our folk had brothers or cousins there and so they’d do. But 
again, that was marrying our relation and no good came of it. I’m related 
to nearly everybody in this place for ten or fifteen miles in every direction. 
Of course we don’t call all of them relatives now in this day. But it is true 





280 


RUTH MOXCEY MARTIN 


that the same blood from various families is in all of our veins and look at 
us. Lots of us up there on your hill (referring to the State Hospital for the 
Insane), lots more of us in the graveyard and we don’t talk of how we came 
there but the town records tell the story, lots more of us (not me especially, 
but men and women living here today) who think perhaps it is the only 
way to do—stop living when you’re better than everybody else on earth.” 
He continued to insist that propinquity, tradition, inertia and in the present 
times decadence and in some cases actual degeneracy all tended to continue 
the pernicious custom of intermarriage, of selection of mates among neigh¬ 
bors’ families, and so among blood relatives. 

In sharp contrast to this town of X-is the town of Y-, in 

another state, yet also seaboard in its location. Tradition says and town 
records, in so far as their incompleteness testify, point to the truth of tradi¬ 
tion, that the first settlers of the prevailing surnames were a group com¬ 
posed of a certain wandering, roistering, loafing, sea-going helper on fishing 
vessels coming from “somewhere south,” his coarse, feeble-minded primi¬ 
tive wife and certain relatives or “pals” of theirs who came with them to 
this port. Some tales have it that this man was an ex-convict. This hermit 
inlet where they could fish, dig clams, get out and in with small boats, 
gather berries inland, loaf generally and still manage to exist suited them 
exactly. To this Lotus-land they gradually attracted people of their own 
sort. Not too many others, for they jealously guarded a spot where they 
could make a living so easily and if too many had come in they might 
have had to bestir themselves—the berry fields would not have been 
inexhaustible. 

One of the field-workers’ chief sources of information was the local station 
agent. Not that the community itself has a railroad. No, indeed, but 
some miles from the dwellings is the stop at which you arrive by train and 
from which you must journey afoot or by conveyance of your own provision 

to reach the actual group of cabins and hovels of Y-. In charge at 

this depot is one of the men who is by blood, birth and residence a man of 

Y-. His only claim to exemption (and it is a great one) is some brains, 

a reasonable honesty and, probably more from his job than through his 
not-to-be coveted heritage, an ability to meet a stranger with a degree of 
frankness. “They are all my kith and kin” he said simply, “but to tell 
you truth if I want to keep anything over night I don’t leave it in my back¬ 
yard nor anywhere but under lock and key in my own house.” This seemed 
to him to stand out as a marked trait of this community composed of his 
kinsfolk. 






INTERMARRIAGE OF BLOOD RELATIVES 


281 


Field work in the town, including calls on a minister and a doctor who 
lived on the outskirts, confirmed this statement. Petty thieving, or better 
stated, an utter disregard of property rights, save out of fear of the owner’s 
fist, was a glaringly evident trait. The clergyman refused to open his mouth. 
He was not of the community he said, but he “hoped to save their souls.” 
He knew what they had done to his predecessor after a write-up of this 
village had appeared in the Sunday morning edition of a large city paper 
—this just after a murder from a drunken brawl in which one of the citizens 

of Y-was prominent. He didn’t care to have the church and his 

dwelling burned to the ground and he didn’t blame the folks for not wishing 
to be made talk of. He saw no point to all this investigation; it was his 
business “to preach on Sunday, marry ’em and bury ’em” and “like Paul 
I know nothing but good of my people”—he was somewhat “set” and the 
field-worker later discovered that his wife was “one of the tribe” though of 
the less undesirable of them. 

The doctor who ministered to their bodies, though seldom, as he said 
“It’s their heads mostly, those can’t be fixed and they don’t care to have 
them,” was communicative. They are as a town, he averred, amazingly 
free (considering their un-morality) from venereal disease. He believed 
that was because they mostly kept to themselves not mingling much even 
with the lowest of towns about. Incest was not unknown; mating of close 
relatives frequent and general, matings “without benefit of clergy” common, 
promiscuity in sex relations before marriage from puberty on very general, 
and shifting about of consorts after marriage not at all astonishing. 

Another physician in a neighboring town informed the field-worker that 

he was sent for in an obstetrical case to call upon Jim-’s wife. He 

went, delivered her of a child. In a year or two he was called to the same 

duty for Dan-’s wife. The woman he found was the former wife to 

Jim. A year or so later he was again called and again for confinement to 
the abode of Jim, where again he found the same woman. This time the 
physician commented and his patient said, “ Well, doctor, I’ve tried ’em 
both and I like Jim the best” and so she lived strictly with Jim from that 
time on. 

A handful of surnames, the same ones, as town records show, from the 
colonial times to the present day have been the chief and almost the sole 
ones of the town. This in itself proves intermarriage has been the rule. 
If anything better has come in, it has been covered and lost. These names 
in larger towns about are almost without fail the most frequent on pauper 
lists. As selectmen in two towns stated “They get in somehow and when 
winter comes they are on our hands for coal and supplies. They are cute, 




282 


RUTH MOXCEY MARTIN 


they try it time and again. If they don’t work it one place or one way they 
do another. They usually take care of themselves summers and then we 
just don’t let them die off winters and that’s what would happen if we drove 
them all back into Y-.” 

Some of the younger ones aged from twenty to thirty do work at times in 
shoe factories in neighboring larger towns. But after thirty years or so 
their manner of life, their feeble-mindedness, their unstable physiques, 
their willingness to “ set about” show. They look like old men and women, 
at least as if they were well on in middle agedness and the younger ones 
bring home to them, if dire necessity drive them to such straits, some 
“cobbling” or certain parts of shoe work which can be done at home with a 
last and hammer from shoe factories of other towns. They have no trolley, 
they trudge in all weathers when trudging must be done; no electricity, no 
water save a few wells and brooks that run to the sea. They never have an 
abundance of food. They seldom have enough. They prefer hunger to 
work and possibly they think little of meal time sav e when hunger calls. 

The circus, the cheap seaside resort with its ferris wheels, its targets for 
prizes, its petty gambling concerns, all are marks for their few pennies or a 
place where they may do some drugdery to pay their way in. As movies 
have come into towns about they do go some to them but always to the 
cheap ones. 

Their sex appetite is appeased from adolesence on and early in life they are 
learned in this lore. They are not especially immoral, but markedly un¬ 
moral. They respond to their sex desires, which are noticeably unsup¬ 
pressed, as naturally and freely as any animal. Physicians who attend them 
grant sex activity is, for the community as a whole, one of their few interests. 
Save for a few who seem exceptionally over-sexed, especially erotic, they 
do not appear to take an indecent attitude toward this part of their lives. 

The mother of twelve pointed to the eldest, a girl of fourteen, and said 
“ she helped me take this last babe at birth. I could have waited on myself 
without even her but I was awful sick and ‘ he’ was away; the doctor I didn’t 
send for, the roads was bad.” The little woman of fourteen was stolidly 
pleased that she had been helpful though she sensed enough of outside 
ways to watch the field-worker closely for any evidence of disapproval. 

Whether or not intermarriage has increased the degeneracy of this stock 
it has kept it true to type through many generations. The pauper list of 
the town has as far back as record goes been almost unbelievable for the 
size of population; in surrounding towns a by-word for generations. 

The town of Z-has far more in common with the first described 

town of X-, though here again is a noticeable difference. Z-’s 






INTERMARRIAGE OF BLOOD RELATIVES 


283 


first settlers were intelligent, probably in many instances something of the 
leader in their make-up. The colonization took place by reason of a church 
dissension. From a town and church some days journey away these pio¬ 
neers who were willing to go through physical hardship and doubtless some 
economic loss for the sake of opinion of conviction, came to a valley (this 
time inland and in yet a third state of New England) to found homes where 
previously only the Indians had lived. 

They were men and women of great religious zeal. They sought to 
convert their Indian neighbors, but were generally unsuccessful. They 
planted churches, one at first which today through sub-divisions of the 
original town has become at least a dozen while the main village of their 
settlement shows four Protestant churches where once was one. Naturally 
in a town of this type there have been frequent instances of pretence and 
hypocrisy but in as many instances have come out of the descendants of 
this settlement men distinguished in theology, in foreign fields of religious 
effort, in pulpits graced by names that are nationally known for sincerity, 
integrity and personality. And from this line of settlers who avowed their 

interest religious rather than as in the case of X -that of family 

property, upbuilding of fortunes by holding of property within the name, 
has come a race of people who have kept almost entirely free from 
intermarriage. 

There are doubtless many reasons for this. While the community has 
through some two hundred to three hundred years been considerably iso¬ 
lated it has still been reasonably “well-known.” It was once offered the 
county seat but its “fathers” sat in council and decided not to let wordli- 
ness and sin creep in. It was once offered the first choice of opportunity to 
a railroad center, but refused. The railroad was put through a small 

hamlet a dozen miles from Z-. Today what manufacturing industry, 

what money to be invested has all been moved to this progressive hamlet 
which is now one of New England’s largest manufacturing cities, while 

Z-is more dead industrially than fifty years after its settlement. The 

marked religious interests of Z-’s people were not of a freak type, but 

of one or two distinctly protestant denominations and their religion and 
ethics undoubtedly stood against intermarriage of blood relatives. Thus 
except for a few of the “black sheep” or a very few instances which any 
group of two to three thousand people through two hundred fifty years can 
show, intermarriage has been practically unknown. It is, however, true 
that mating has depended much upon propinquity further limited by iso¬ 
lation of community, until though actual relation in marriage is rare, the 
town is an almost inextricable group of “by marriage” relatives. This 






284 


RUTH MOXCEY MARTIN 


coupled with a large influx of undesirables soon after the Civil War and by 
a two hundred and fifty years continual exodus of the more progressive, and 

a vegetation right in Z-of those too inert or inapt to make their way 

in the great west, has worked for a very evident decadence which is at many 
points actual degeneracy. 

In summary it is doubtless true that X-with its intelligence and 

thrift is showing more and more a tendency, in this day of breaking from 
tradition, to out marriage. It will mean in some instances eugenic salva¬ 
tion; in others where the mental taint has become very marked it will but 
serve to carry into other strains of Yankee blood the cacogenic factor 
resulting most often in X-in insanity and suicide. 

No doubt Y-is hopeless. It will in time accomplish its own ex¬ 

termination yet the town is a menace to the country for it is true that an¬ 
nually a few of the “prettier” girls or the less feeble-minded boys do drift 
out into the surrounding towns and marry. Wherever they go their traits 
hold true and the menace is especially great in instances where a reasonably 

eugenicNew England man sets aside his feeling about Y-and marries 

a pretty Y-girl. This girl practically always bears the man a large 

family of children and in few instances are they other than true descendants 
of Y-, feeble mentally, moron at the most. 

Z’s probable uneugenic future is as indicated not to come from intermar¬ 
riage, but rather from the fact that into her valley will continue to drift 
the unambitious, the uneugenic, those who prefer to avoid law, order and 
effort in the world outside—while her better stock will as it has for nearly 
three centuries tended to do, go out a few at a time in the persons of the 
youth of the town for greater opportunity. 









FORMAL INBREEDING IN HUMAN SOCIETY WITH SOUTH 
AMERICAN EXAMPLES 


HERBERT J. SPINDEN 
Peabody Museum of Harvard University 


The simple biological family, that is, the group composed of mother, 
father, and children, is universal and psychologically important among 
all human beings—normally speaking—while regulations in marriage 
carrying us at least a step beyond the natural family, are constantly seen in 
the larger organizations of human society. A fundamental classification 
of the social structures developing out of the simple family may take the 
following form: 


(UNIVERSAL AND FUNDAMENTAL BUT NON-CONTINUQUS ) 


CLOSED ONE-SIDED FAMILY 
(FORMAL EXOGAMY) 


CLOSED TWO-SIDED FAMILY 
(FORMAL ENDOGAMY) 


OPEN TWO-SIDED FAMILY 
(FORMAL EXOGAMY) 


CLAN 

GENS 

CROSS-COUSIN 

BROTHER-SISTER 

(ELIMINATION 

OF FATHER’S 

(ELIMINATION 

OF MOTHER’S 

MARRIAGE 

MARRIAGE 

SIDE) 

SIDE ) 

UNCLE-NIECE, 

AUNT-NEPHEW 

J 

1 

| MARRIAGE J 


Fig. 1 


Natural or Biological Family 


Social classes, aristocracies, royal families, theocracies, etc., may develop 
out of any of these bases: 

The Race (Common Blood) 

I 

The Stock (Common Language) 

I 

The Nation (Common Culture) 

The Lateral Development of the Family 
Pattern of Cross-Cousin Marriage on the Field of Consanguinity 
Diagram of Uncle-Niece, Aunt-Nephew Marriage 
Pattern of Clan System on Field of Consanguinity 
285 













286 


HERBERT J. SPINDEN 


Inheritance is on the mother’s side, solid black. Father’s side, shaded, is not repre¬ 
sented among his own children, but instead among his sisters’ children. The children of 
both the mother’s brothers and the father’s brothers belong to the clans of their mothers, 
C, Cl; D, Dl, etc. Finally, E, El; F, FI, etc. record possible contacts with other clans 
established through marriage since the husbands of the maternal and paternal aunts 
must come from outside the clans of these women. Clan A may be substituted for D, D1; 
F and FI, and Clan B for C, Cl; E and El reducing the exogamic groups to two. 

The consequences of regulated inbreeding among human beings have a 
very real interest for students of eugenics. These students cannot expect to 
observe many cases of consecutive human inbreeding in the course of their 
own lives and they must either turn to the past or content themselves with 
applying to man certain principles of inheritance induced from observa¬ 
tions on plants and animals with much shorter generations. If they turn 
to the past they often find insuperable difficulties in the matter of record. 
In certain favored localities, when genealogies are sufficiently detailed, 
inbreeding may take place but rarely. 

I wish, therefore, to call attention to localities and peoples where there is 
reason to believe that a large percentage of the marriages have been of the 
endogamic or inbreeding type and where effective inbreeding may have been 
the rule over long stretches of time. It should be possible for properly 
qualified investigators to observe in some of these localities the results of 
close marriage in three or four generations of living individuals and to 
obtain information from older persons going back two or three generations 
into the past. I can present no data on the physical characters which might 
seem to have been determined or emphasized by such inbreeding, merely 
indicating fields where future eugenic research should prove profitable. 

What I have here called the open two-sided family is perhaps the com¬ 
monest form. As a rule the mother’s side and the father’s side are equally 
important, or, at least, both are held in consideration. Here we find a 
rigid prohibition, on moral grounds, of marriage within the closer degrees 
of consanguinity on both sides of the house. Partial inbreeding may result 
from consanguinous marriages which approach the prohibited degrees, 
especially in wealthy, aristocratic or ruling families. Among some southern 
European peoples marriages between cousins, uncle and niece, etc., are 
common, but never the rule. The normal human family is not a profitable 
field in which to study inbreeding. 

The one-sided family (the term one-sided referring only to extensions 
beyond the simple family) occurs in both hemispheres and exhibits some 
very interesting convergencies. Anthropologists of the United States 
have come to a fair degree of accord as regards the words “clan” and 
“gens,” the first referring to social groups in a tribe or village that follow 


FORMAL INBREEDING IN HUMAN SOCIETY 


287 


the mother-line and the second to similar groups that follow the father-line. 
Both are normally exogamic. There are often larger groups, made up of 
several clans or gens, and as regards these larger groups (phratries, if several 
in the tribe, or moieties, if only two) marriage is sometimes regularly inside 
the group and sometimes regularly outside. Theoretically neither the 
clan nor the gens is regarded as an extended one-sided family although often 
explained as such. The possibility is always held that a clan or gens may 
contain two or more families. The clan or gens is chiefly important because 
it serves as a basis of organization in religious or civil government. It 
ordinarily takes its name from, first, a tutelary being or totem; second, some 
object of nature which may or may not have a ceremonial relation to the 
group; and third, some geographical location which may, perhaps, be con¬ 
sidered as the original home of the group. Clans and gens in different parts 
of the world are often superficially similar, while inwardly diverse. In 
relation to problems of breeding they are not nearly so important as often 
assumed to be. 

Real inbreeding can only take place in a closed circuit which must take 
account of the inheritance from both parents. In the closed two-sided 
family marriage must ordinarily be held within a limited field of consan¬ 
guinity. In practice, among so-called primitive people (who have just as 
long an ancestry as civilized people), the commonest form of close marriage 
is cross-cousin marriage. Of course any form of cousin marriage would be 
just as effective as this, for breeding purposes, but it happens that marriage 
between cousins who are the children of fathers’ sisters or mothers’ brothers 
is much more frequently dictated by custom than that between parallel 
cousins (children of mothers’ sisters or fathers’ brothers) or simply un¬ 
differentiated cousins . 1 With a difference of one generation, marriage is 
sometimes also formally demanded between nieces and their maternal 
uncles and between nephews and their paternal aunts. These two types of 
marriage prevailed among nearly all the important tribes and linguistic 
stocks of South America. Brother and sister marriage was formally 

1 Tylor in 1889 first called attention to the institution of cross-cousin marriage. He 
says: ‘‘In tabulating the nations of the world, I found a group of twenty-one peoples whose 
customs as to the marriage of first cousins seemed remarkable; it is that the children of 
two brothers may not marry, nor the children of two sisters, but the child of the brother 
may marry the child of the sister. It seems obvious that this ‘cross-cousin marriage’ 
as it may be called, must be the direct result of the simplest form of exogamy, where a 
population is divided into two classes or sections with the law that a man who belongs 
to Class A can only take a wife of Class B.” (On a method of investigating the develop¬ 
ment of institutions; applied to laws of marriage and descent, Journ. Anth. Inst., XVIII, 
pp. 245-269.) 


288 


HERBERT J. SPINDEN 


demanded among the ruling Inca of Peru. It probably does not survive 
today among any people except, perhaps, as an occasional anomaly. The 
levirate, where a man takes his dead brother’s wives, and the somewhat 
analogous situation where sisters become the polygamous wives of the same 
man, or succeed each other in this service, are probably responsible for the 
development of the cross-cousin relationship as we shall see presently in a 
diagram. Both of these usages prevail over large parts of South America. 

Now, I am aware that the cross-cousin relationship is recognized in the 
name when it has no present part in actual marriage. Among many 
American Indian tribes, with or without the clan or gens system, the chil¬ 
dren of mothers’ sisters and fathers’ brothers, that is the parallel cousins, 
are called by the same terms that signify brother and sister while the cross¬ 
cousins are designated by another term. 

The classical comparison of Morgan between the relationship terms of 
the Iroquois of North America and the Tamil of southern India shows that 
the cross-cousin relationship complex was completely expressed in both. 
But in the case of the Iroquois, at least, there was no cross-cousin marriage. 
Simply, the term “father” covered a group including the real father and 
his brothers and the term “mother” covered a group including the real 
mothers and her sisters. The children of these various persons (and of 
outside persons joined to them in marriage) regarded each other as brothers 
and sisters. On the other hand, the sisters of the father were aunts, the 
brothers of the mother, uncles, and their children were grouped together as 
the only cousins. 

Marriage among the Iroquois was exogamic as regards the social groups 
known as clans, and the membership in a clan was inherited through the 
mother. The real consanguinity system and the conventional clan system 
did not coincide. The group of persons that an individual could call his 
brothers and sisters was divided among several clans. The real brothers and 
sisters, plus the so-called brothers and sisters who were children of the 
mother’s sisters, belonged to the mother’s clan. The so-called brothers 
and sisters who were children of the father’s brothers belonged to the 
various clans of their own mothers who had married these men. Similarly 
the cross-cousins, although grouped under one name, represented different 
clans. The children of the maternal uncles belonged to the various clans 
of the wives of these men, while the children of the paternal aunts belonged 
to the clan of the real father and his brothers, inherited from their common 
mother. 

The explanation might be carried farther but enough has been said to 
show that clan-grouping and essential relationship grouping did not coin- 


FORMAL INBREEDING IN HUMAN SOCIETY 


289 


cide among the Iroquois. Nor do they coincide among other tribes having 
the clan or gens system. It is not enough to say of any tribe that marriage 
is exogamic as regards the clan or gens. Marriage is also controlled by 
consanguinity at one and the same time that it is controlled by the clan or 
gentile organization. The Iroquois could not marry those members of the 
brother-sister group who were outside their own clan. It is doubtful if 
they could marry their father’s brothers and sisters—that is, their paternal 
aunts and uncles—although these persons were also outside of their own 
clan. Whether or not they could marry the cross-cousins, who were all 
outside their clan I am not able to say but among some tribes with clan or 
gentile organization the marriage of cousins is forbidden, or frowned upon. 

The statement so often made that the clan or gens is an effective device 
to prevent inbreeding is utterly fallacious. It does of course prevent the 
marriage of certain relatives but it does not, by itself, prevent union of 
the same blood. It would be theoretically possible to have in a full-blown 
exogamic clan or gentile system a very effective kind of inbreeding. This 
type of social organization neglects one half of the extended family, as has 
already been pointed out. In other words, the barriers against inbreeding 
affect only one half of the field of consanguinity. Inbreeding results when 
the same blood is turned back continuously into the stream of descent 
through marriage within the real family. The clan or gentile system is a 
social superstructure that can be entirely neglected by students interested 
in inbreeding the races and strains of the human species. 

If inbreeding is possible in types of human society which have apparently 
raised up barriers against it, and yet does not occur in the normal human 
family or at least does not occur with sufficient regularity to have a deter¬ 
minable eugenic effect, what does constitute good evidence of actual 
inbreeding? Let us examine a typical field of consanguinity covering two 
generations, giving all the type individuals. The same blood (this term 
must be taken to mean the complex of physically heritable characters) 
is present in all these type individuals possibly weakened in the collateral 
lines by the unrecorded second parents of the cousins. 

The relationships given here are identical with those in the diagrams that 
follow. 

Since any table of consanguinity must be made formal for comparisons 
it is understood that the three children of the mother and father (namely, 
Ego and this brother and sister), here shown on the top of the diagram, are 
reversible to the bottom of the diagram to complete the father line. It is 
assumed here that each parent has two brothers and two sisters and that 
each of these has two offspring, one male and one female. The husbands 
and wives of the collateral aunts and uncles do not appear in the diagram. 


290 


HERBERT J. SPINDEN 















FORMAL INBREEDING IN HUMAN SOCIETY 


291 


Now if we superimpose upon this field of consanguinity the relationships 
established by the levirate, whereby the husband’s brothers may succeed 
to his marital rights and the reciprocal system for which the term sororate 
has been suggested, whereby the sisters of the wife may assume her duties 
either by polygamous marriage or by succession we have the following 
aspect of the field of consanguinity: 


BROTHERS AND SISTERS COUSINS 



COUSINS BROTHERS AND SISTERS 


(FATHER LINE AFFECTED BY THE LEVIRATE,MOTHER 
LINE BY INCLUSION OF HER SISTERS) 

Fig. 3 

Most convergences in human culture are brought about through some 
mechanistic constant. Given the natural family and the cooperative in¬ 
stinct, the assumption of the duties of a deceased husband by his brothers, 
and of a deceased wife by her sisters, is one of the few methods by which 
continuity of the group can be accomplished. It appears to me that the 
wide but broken distribution of the cross-cousin relationship complex is 
explained by the formal, or mechanistic, answer to this common problem 
of continuing the family organization contained in the above diagram. 

The cross-cousin relationship is found among many tribes where cross¬ 
cousin marriage does not prevail, as has already been stated. It need not 
be assumed that cross-cousin marriage ever existed among such tribes, 
since the primary obligations of parental protection are sufficient to account 
for the form of organization. Farrand found the cross-cousin relationship 
perfectly developed among the Alsea of Oregon although the prevailing 
type of marriage was outside the tribe. But given the fundamental 
arrangement, then cross-cousin, uncle-niece, and aunt-nephew marriage 
might easily develop. 

Any male of the brother-sister group, solid black, can marry any female 
of the cousin group, shaded, either laterally or across the field, and vi^e 


versa. 




292 


HERBERT J. SPINDEN 


BROTHERS AND SISTERS COUSINS 

MALE FEMALE MALE FEMALE MALE EGO FEMALE MALE FEMALE MALE FEMALE 



MALE FEMALE MALE FEMALE MALE FEMALE MALE FEMALE 

Fig. 4 


The uncle-niece, aunt-nephew marriage presents an offset in the genera¬ 
tion, so to speak. It doubtless occurs in combination with cross-cousin 
marriage rather than as a complete substitute for it. With this type of 
marriage the board is not cleared in two generations. Indeed it seems 
impossible that a marriage system could continue as a going concern where 
young men marry old women and old men young women unless each should 
mate once in youth and again in middle or old age. The diagram of uncle- 
niece and aunt-nephew marriage follows: 


NIECE NIECE NEPHEW NIECE THIRD GENERATION INBRED 




Q 0. 0 O 


MATERNAL UNCLES' 




PATERNAL AUNTS 

P O O O 

THIRD GENERATION INBRED 



NEPHEW 



o 


NEPHEW 


Fig. 5 


It will be observed that all the members of the extended brother and sister 
groups of the previous tables do not analyze into nieces of the father’s 
sisters and nephews of the mother’s brothers. And the offspring of these 
marriages of different generations (in the position of the earlier cross¬ 
cousins) properly belongs to a third generation. It would seem that the 
cross-cousin marriage was a more perfectly operating system than uncle- 
niece or aunt-nephew marriage, but the two could probably be used in 
conjunction and the evidence shows that they were so used in some Sbuth 
American tribes. 

















CUN A - MOTHERS CLAN I MATERNAL UNCLES’wiVES’ CLANS 



I 




























294 


HERBERT J. SPINDEN 


Let us now look at the same field of consanguinity when a formal clan 
system has been laid down upon it. Here we find it necessary to take 
account of affinity as well as of consanguinity since the clan inheritance 
follows the female who must always be of a different clan than the male 
with whom she mates. We have already seen, however, that a close type 
of inbreeding may take place in spite of clan and gentile restrictions. 

If we had to depend entirely upon the direct statements in ethnological 
studies concerning close marriages in South America we would not consider 
our position strong. There are some clear references but they do not begin 
to cover the field. Thus Gilij speaking of the Orinocan Indians says: 

. . . . Many among the Tamanchi, select for wives their nieces, daughters of 
sisters. The maternal uncle is there called by the same name as otherwise is given to the 
father-in-law. 2 .... But there are no cases in which a young girl takes for a hus¬ 
band her paternal uncle. And after much consideration I have found that this is because 
they have the habit of calling him father. 

Jean de Lery writes of the Indians of Brazil in the sixteenth century: 

Touching the marriage of our Americans, they observe only these three degrees of 
consanguinity; namely no one takes his mother, his sister or his daughter to wife, but 
as to the uncle he takes his niece and furthermore in all the other degrees of relationship 
no regard is paid. 

The fact of uncle-niece and aunt-nephew marriage caught the attention 
because of the wide differences in age but it was not often grasped that the 
uncles were mother’s brothers and the aunts were father’s sisters. The 
mating of cousins is also noted by various early writers particularly among 
Carib and Arawak tribes. Thus Barrere, referring to the Galibes and other 
Carib tribes of French Guiana says: “Ordinarily the Indians do not make 
misalliances. Always they espouse their relations, even to the second degree 
of consanguinity. The youths regard their blood cousins as belonging to 
them by a certain right of birth. So they often marry them, even when 
they are only two or three years old. In the mean time another woman is 
taken who is sent away when the young cousin has grown sufficiently to 
enter into relations with her husband.” 

Priests are frequently outraged at the close marriages and assert that 
nothing except the first degree of consanguinity was sacred. On the other 
hand Roth tells us that the Arawak sometimes accused the Spaniards of 
incest. This was when they married parallel cousins instead of cross¬ 
cousins. Recently Senor Lefone Quevedo described the close-marriage 

2 This statement indicates that cross-cousin marriage also obtained: only in this way 
can the maternal uncle become the father-in-law. 



FORMAL INBREEDING IN HUMAN SOCIETY 


295 


customs of the southern Guarani. The present writer read a paper on the 
social organization of the Mosquito Indians showing that sexual hospitality 
is still the rule between unmarried cross-cousins and that normal marriage 
is within this relationship. 

There is a kind of evidence which is abundant, and I believe conclusive, 
namely linguistic information. The Catholic Church forbids certain 
consanguinous marriages and permits others only after the payment of a 
fee for dispensation. As a result of this regulation we find a list of relation¬ 
ship terms in nearly every grammar and catechism written in South and 
Central American languages as well as in the occasional dictionaries. 
At first glance it might not seem that practically conclusive evidence of 
inbreeding customs could come from such sources. 

But think of the coincident relationship terms when real relatives ac¬ 
quire a second title through marriage! With cross-cousins mating the 
following relationships coincide. 

1. Cousin = husband = wife 

2. Cousin = brother-in-law = sister-in-law 

3. Maternal uncle = father-in-law 

4. Paternal aunt = mother-in-law 

5. Nephew = son-in-law 

6. Niece = daughter-in-law 

When, therefore, you find the same word having a double meaning which 
could only come about through cross-cousin marriage it is a pretty safe 
guess that this form of marriage existed. Just as an example: we know 
little enough about the Mosetenes Indians of Bolivia but in a short word 
list we find that vi means either brother-in-law or first cousin male, while 
font means either sister-in-law or first cousin female. Such a coincidence 
of terms could only be effected through cross-cousin marriages. I have not 
space to present the linguistic data which are rather voluminous, but I 
may say that among nearly all Arawak, Carib, Tupi, Guarani, and Chibcha 
tribes as well as a number of smaller South American tribes the linguistic 
evidence of this double meaning of relationship terms in consanguinity and 
in affinity is mute testimony of a custom of inbreeding among these Indians. 

Clan-like structures (except that they are probably endogamic family 
groups instead of exogamic groups) are sometimes found, as among the 
Bri bri of Costa Rica, the Goajiro of Colombia and Venezuela, the Araucano 
of Chile and possibly the Inca of Peru. Let us examine briefly the affairs 
among the last mentioned people. In the preamble of Incan history the four 
Children of the Son, with their wives, lead ten ayllu, or lineages, out of 
Tampu Focco, and, after a peregrination in which three of leaders were 


296 


HERBERT J. SPINDEN 


killed or turned into stone idols, the ten ayllu settled at Cuzeo. The ayllu, 
in origin at least, was probably expanded endogamic families. According 
to Markham the marriage of brothers and sisters was a late development of 
the Inca royal house. This custom is also reported for the Nachez tribe of 
the lower Mississippi where the rulers, as in Peru, were regarded as children 
of the Sun and therefore divine. It is possible that brother and sister-mar¬ 
riage was made necessary by there being no other person of equal rank for 
each to marry. But surely such a custom would be more likely to grow out 
of endogamic practices such as cross-cousin marriage, than out of exogamic 
clans or gentes. Senor Lef one Quevedo in a letter to me containing valuable 
notes on South American relationship terms considers that the Peruvians 
had clans—but he would doubtless agree that these clans were endogamic 
rather than exogamic and, by definition clans must be exogamic. 

Among other tribes the social structure was by villages. Farabee notes 
among the Arawak tribes of the southern British Guiana region cross¬ 
cousin marriage, that is outside the village. Thus family A is distributed 
in two or more villages, the girl must marry out of her village but within her 
family. 

In closing, I only wish to emphasize the fact that close marriage was the 
rule rather than the exception among many South American and some 
Central American tribes and that it is still maintained. Of course it is not 
to be expected that any type of marriage could be maintained in pure form: 
love laughs at locksmiths. But enough intermarriage might have taken 
place to intensify family types among these tribes. Through its eugenic 
significance somatoJogical research in South America should take on a new 
impetus. Skeletal work might also be carried out with the new thought in 
mind. It would seem that South America was the most promising field in 
which to study the results of human inbreeding. 


THE EFFECT ON THE GERM PLASM OF ISOLATION IN A 
MOUNTAIN SECTION 

W. S. ANDERSON 
University of Kentucky 

INTRODUCTION 

One of the bluegrass counties of Kentucky extends to the foothills of the 
mountain section of the state. There is a strip of territory in this county 
four or five miles wide and eight or ten miles long just at the foot of the hills. 
This strip of land is rolling and traversed by small streams. The streams 
attain considerable volume during rainy seasons and some of them cease 
to flow at all during the dry summer months. 

While most of the county is a very fertile and prosperous farming section 
of Kentucky the plateau at the foothills is a sandy clay almost worthless 
for the purpose of cultivation. The bluegrass land of the county is eagerly 
sought by farmers at from $150 to $300 per acre while this plateau soil can 
be purchased at from $5 to $40 per acre. 

There is no natural barrier of any kind between the two classes of soils. 

At a certain very irregular line the rich phosphatic limestone disappears 
to lower depths and a shale, or sandstpne crops out. The character of the 
two soils is as dissimilar as the strata out of which they have been formed. 
The one will grow bluegrass, clover, corn, wheat, hemp, tobacco, etc; the 
other is cultivated almost exclusively in corn and the return per acre is 
only a fraction of what good land produces. 

ORIGINAL SETTLERS 

About ninety years ago I J. M. and wife settled on this plateau. No 
information can be procured concerning their ancestry. The wife, as 
far as records show, was of good physique but of less than average intellect. 
I J. M. is described by a grandson, III A.M., as a man of very high temper 
which he was unable to, or at least did not, control. He had chorea. There 
are no records to show when the involuntary movements began in his case, 
but at the time he is remembered by III A. M. he had difficulty in feeding 
himself, requiring both hands instead of one to carry food or liquid to his 
mouth. 


297 


298 


W. S. ANDERSON 


All the children of this couple drifted away from the plateau except one 
son, II E. M. The descendants through his sons and daughters who left 
this section of the state have been traced but no cases of chorea have been 
found among them; nor are they noted for any antisocial traits. We are 
only concerned with the descendants of the son who remained at the place 
of his birth. 

The son, II E. M., who remained at home was affected with chorea in 
his old age much as his father had been. He was devoid of ambition and 
unable to obtain by his labor more than a bare living for himself and family. 
He was content to live in a log cabin, cultivate a few acres of unproductive 
land in corn and hunt and fish for his meat. 

About the same time that I J. M. came to the plateau I A. W. took 
possession of another part of it. I A. W. and wife were quiet, unambitious, 
and mentally deficient, perhaps they should be classed as high grade 
morons. They were strong of body and able to do a great amount of 
physical labor. They did not work, however, except under the spur of 
necessity. 

For a home they built a log house, covered with clapboards held in place 
by poles. The chinks between the logs of the house were partly filled with 
clay mortar. All furniture was made out of rough timber. If there was 
food for one day little thought was given to the needs of other days. Each 
spring a small plot of the unproductive soil was planted in corn. 

To I A. W. and wife were born a large number of children, some ten or 
twelve living to maturity. The children, having high grade feeble-minded 
parents were all of the moron type, none of them showing more industry, 
ambition or intelligence than their parents. 

A daughter, II B. W., of I A. W. married II E. M. the choreic son of 
I J. M. 

In this marriage there is the union of two streams of germ plasm both of 
which are characterized by feeble-mindedness, the family of the husband 
carrying the taint of chorea as well as the moron tendency. 

THE DESCENDANTS 

To IIE. M. and IIB. W. were born four sons and three daughters. The 
descendants of the sons and daughters of II E. M. and II B. W. now con¬ 
stitute the entire population of the plateau and consist of 150 families 
numbering 900 people. The present population is as deeply inbred as a 
population can be not to allow brother and sister marriages. Cousin 
marriages and double first cousin marriages are common. Some blood 


EFFECT ON GERM PLASM OF ISOLATION 


299 


outside the two original families at first had to be introduced but now there 
are few or no marriages that do not trace on one or both sides to the one 
or the other foundation family and frequently to both. 

MAKING PURE THE GERM PLASM 

Had a breeder of domestic animals set out to produce an extensive herd 
of cattle all of which would breed true for two features, one a dominant and 
the other a recessive, he would have adopted by selection the exact method 
in making his matings which these people used in their marriages. The 
breeder would have striven to get his germ plasm pure for both traits. 
That is what has happened to the human germ plasm which has produced 
the plateau people. Feeble-mindedness is recessive. The original families 
both were pure for the traits and their children were pure. It has so hap¬ 
pened that no out crosses have been made with normals, but always with 
subnormals and this has kept the stream pure. So far, then, as a deficient 
mental makeup is concerned the stream is pure and the fourth and now the 
fifth generation is breeding true. 

The other trait, Huntington’s chorea, is dominant and originally was in 
but one of the foundation families. By continual intermarriages of close 
relatives it has now become so near universal that some members of every 
family are victims and in many families all members suffer from some form 
of it. The cattle breeder to have introduced a dominant trait would have 
in his wisdom made the same close matings which the isolation and igno¬ 
rance of these people have caused them to make. 

♦ 

THE EXTENT OF THE TRAITS 

The writer can think of no hereditary human trait that is less desirable 
than chorea or one that in marriage should be more carefully avoided; yet 
here is a community that has fostered the “serpent” until every family is 
cursed by it. The germ plasm, by ignorant, consanguineous marriages, 
is rapidly becoming pure for chorea as it is for feeble-mindedness. 

From the original progenitor, I J. M., there are now hundreds who carry 
and can transmit the trait; in fact, are transmitting it. 

The two factors which have been most influential in tainting the stream 
of germ plasm are the low mentality of the population and the segregation 
of the people. 

Were the people of normal intellectual endowment their knowledge 
would prevent the young people from marrying into the tainted families. 
Were they stronger mentally they would not remain in the community 


300 


W. S. ANDERSON 


which offers no opportunity for any of the comforts or pleasures of life. 
They would go where homes, and schools, and churches, and soils offer 
some commensurate return for energy expended. No natural barrier 
separates them from a most prosperous territory. The reason they stay 
on the poor soil is their lack of ambition, their absence of initiative and their 
willingness to congregate together. 

It causes no very great concern to find one or two choreics in a community, 
but it is alarming that every home in a given territory has its victims. In 
fact it becomes a nuisance to the county and even to the State. 

Owing to the primitive conditions under which the people live few of them 
have been sent away from home for care and treatment. Cases of extreme 
violence are kept at home by the neighbors coming in to aid in restraining 
them, which is done by physical force. A lack of refinement, and igno¬ 
rance of the equipment and care given patients in modern asylums cause 
them to keep in their homes choreic patients who have reached the irre¬ 
sponsible mental stage in addition to their lack of muscular control. 

BIOTYPES OF CHOREA 

Davenport and Muncey 1 showed that Huntington’s chorea has many 
species or biotypes. The chorea of the plateau manifests itself in many 
different ways in the different families. The age of onset varies. In some 
families the onset is after the age of menopause, while in others it is near the 
age of physical maturity and in still other families it manifests itself before 
the age of twelve. A few cases, as a seeming result of double cousin 
marriages, are recorded in which the infants were afflicted from birth. 

One type manifests itself in very violent muscular movements especially 
if the patients are young. The name given to the disorder by the com¬ 
munity is the “jerks.” Strong men are required to hold children ten and 
twelve years of age in bed, when the attacks are at their most violent stage. 
In this severe type mental irresponsibility continues as long as the muscular 
contraction and improves as the treatment relieves the involuntary 
movements. 

Another type is represented by V. Y. M., a young married woman. 
There was little involuntary movement in her case; but temporarily there 
was manic-depressive insanity. Her family would not consent to send her 
to the asylum and she constantly tried to commit suicide by drowning 
although there was no water nearby save a small stock pool. She would 
throw herself into this if left free for a few minutes. 

1 1916 Bulletin No. 17, Eugenics Record Office. 


EFFECT ON GERM PLASM OF ISOLATION 


301 


One of the alarming types makes itself manifest at an early age, especially 
if there be any unusual stress. A representative example of this is the case 
of V. M. M., fourteen years of age. In June, 1921, she married a cousin 
sixteen years old. Immediately after assuming the marriage relation chorea 
developed in such a severe form that she lost all control of her limbs and 
voice. Separation from the husband and treatment are gradually restoring 
her health; but restoration is doubtless temporary as the universal rule for 
this type which appears so early is to recur again and again. 

Perhaps, the most common type of the chorea manifests itself in the 
uncontrollable tempers of the victims. They are not mere irritations but 
intense outbursts of anger that lead to deeds of violence and murder even 
if the object of the anger is of the same family. Before and after the storms 
of temper there are the tremors and twitchings of the muscles which the 
patients cannot control any more than they seem to be able to control 
their angry passions. 

The original carrier of the choreic germ plasm, I J. M., was noted for his 
violent outbursts of temper. It seems that in going on from one generation 
to another this disposition has become intensified until it is difficult to give 
in words the hold which it has upon his descendants. Even those who have 
little or no physical manifestations of chorea are characterized by un¬ 
governable tempers. 


THE MORALS OF THE PEOPLE 

In the territory described there is segregated a considerable population 
rapidly degenerating to a pure race of choreics and imbeciles. Already the 
community is noted for its lack of a wholesome moral sense. At one time 
the men were skilled woodmen. They had a steady income from the sale 
of railroad ties, mortised fence posts, and logs ready for the saw mill. Long 
since the timber has been exhausted. The present generation exists on as 
little work as possible. 

They consider it legitimate to sell their vote and this is freely practiced 
by both men and women. The distillation and sale of white whiskey 
promises to be their main industry. The low moral sense is severely con¬ 
demned by the adjoining communities but the condemnation is made 
because the reason is not fully understood. The writer believes it is as 
natural as the uniform physical appearance of blue eyes and light hair 
and certain facial features. 

We wish to make it clear that the isolation of the plateau is due to the 
lack of ambition in these people. They live under the hard conditions 
because they want to. They know that they do not have ability and ambi- 


302 


W. S. ANDERSON 


tion to go to the Bluegrass country nearby and set up a home. They are 
poor and they are making no effort to be otherwise. They love their kind. 
The sex appetite is their ruling passion but they believe it should be con¬ 
trolled until marriage. The result is that they often marry just at the end 
of childhood, long before they are fit either physically or financially. In 
fact they seem not to consider money or property necessary for marriage. 
If there is the impulse the marriage takes place and two immature people 
start a family of their own although the only prospective income is the 
uncertain day wage of the boy. The size of the family is only limited by 
the inability of the wife to reproduce and ranges from six to fifteen children. 
The ignorance of the mothers and their lack of sanitary conveniences cause 
a very high infantile death rate. 

The in-and-in marriages, due in part to the voluntary isolation and in 
part to the inability of the young people to find marriage partners outside 
their own relatives, have resulted in concentrating in their germ plasm the 
factors for chorea, ungovernable tempers, imbecility, indolence, and a 
general anti-social disposition. 


DfiVELOPPEMENT COMPARE DES PRODUCTS SUCCESSIFS 
D’UN MfiME COUPLE 

ETIENNE RABAUD 1 
Paris , France 

Suivant une croyance populaire fort repandue le premier ne d’un 
couple serait moins bien doue sous divers rapports que ses freres et soeurs 
puines. Cette croyance, comme toutes les autres, ne reposent evidemment 
que sur des faits mal observes ou sur des prejuges sans fondement; elle 
s’impose neanmoins a bien des esprits comme une verite demontree. Ceux-ci 
en tirent alors la conclusion que dans un pays oii les naissances sont peu 
nombreuses les aines constituent une grande partie de la population qui, 
de ce fait, se trouve composee d ’individus plus ou moins tares. 

Divers evenements, la guerre en particulier, sont pour cette croyance 
un veritable controle en donnant aux diverses qualites individuelles l’occa- 
sion de se manifester. II n’est pas inutile cependant de proceder a un 
examen rigoureux. Des statistiques bien conduites sur la descendance 
d’un tres grand nombre de couples humains fourniraient un resultat 
decisif. Ces statistiques n’existent pas et semblent peu faciles a etablir. 
C’est pourquoi j’ai pense qu’en etudiant la descendance d’un certain 
nombre de couples, d’une espece animale quelconque on obtiendrait des 
indications suflisantes. Je rapporte ici les resultats de mes recherches sur 
les souris, me reservant d’en donner ulterieurement un expose detaille. 

En these generale, je puis affirmer que mes recherches ne confirment 
nullement la croyance populaire. J’ai organise mes couples de fagon 
a repondre autant que possible a toutes les conditions: parents jeunes, 
parents vieux, parents d’ages tres differents, l’un jeune, 1’autre vieux, 
parents consanguins, parents etrangers l’un a 1’autre. Aucune des condi¬ 
tions etablies n’a mis en valeur de differences appreciates; ni les unes ni les 
autres ne determinent chez les individus de la premiere portee des qualites 
physiques inferieures a celles des individus des portees suivantes. Chaque 
portee etait pesee a la naissance, puis regulierement pendant quatre se- 
maines consecutives et une derniere fois au bout du troisieme mois.. Des 
pesees supplementaires etaient faites quand le besoin s’enfaisait sentir. 

1 Pr6sente par M. Marin Molliard. 

303 


304 


ETIENNE RABAUD 


Dans un grand nombre de cas, les portees successives d ’un meme couple 
se comportent de la meme maniere et leur croissance suit une marche 
parallele. Voici, a titre d’exemple, les poids moyens relatifs aux cinq 
portees d’un couple: 



1 ERE PORTAE 

2 E1TE PORTAE 

3 EME PORTAE 

4 feME PORTAE 

5 EME PORTEE 


Petits 

Grammes 

Petits 

Grammes 

Petits 

Grammes 

Petits 

Grammes 

Petits 

Grammes 

1 ere pesee ... 

10 

1.25 

10 

1.45 

5 

1.25 

4 

1.56 

5 

1.05 

2 eme pes6e... 

8 

2.60 

8 

2.73 

5 

2.05 

4 

3.56 

3 

2.66 

3 eme pesee... 

8 

3.87 

7 

3.71 

5 

3.95 

4 

4.57 

3 

4.08 

4 eme pes6e... 

8 

4.78 

7 

4.42 

5 

5.85 

4 

6.81 

1 

6.00 

5 eme pes6e... 

8 

7.85 

7 

6 

5 

9 

4 

11.12 

1 

8 

6 eme pes6e... 

8 

25 

7 

23 

5 

22 

4 

24 

1 

24 


D’une portee a l’autre existent evidemment des differences; mais 
elles sont insignifiantes et le resultat final est en somme le meme. On 
remarquera au surplus que le nombre des petits allaites n’influe pas 
sensiblement sur la croissance. 

II est evidemment des cas ou les differences entre les portees sont plus 
accusees; elles sont alors tan tot en faveur de la premiere portee, tantot en 
faveur de la seconde. Parfois, il arrive que, la deuxieme portee se com- 
portant mieux que la premiere, la troisieme se comporte moins bien. Tel 
est le cas de mon couple 57 dont les produits de 1 ere et 2 erne portees pesent 
respectivement 20.50 et 22.50 grammes au troisieme mois, tandis que ceux 
de la troisieme portee pesent seulement 13 grammes au bout du meme laps 
de temps. 

Dans ce cas, comme dans d’autres semblables, je n’ai pas observe de 
maladie intercurrente. Mais, en depit de toutes les precautions prises, 
l’experimentateur n’est pas maitre de toutes les variables ni toujours en 
etat de les connaitre. On ne peut pourtant attribuer des differences de 
l’ordre de celles que je signale qu’a des modifications peut-etre passageres 
survenues dans l’etat des parents. 

D’une fagon tres generate, je retire de mes elevages l’opinion que la 
croissance des descendants d’un couple, dans les conditions normales de 
nutrition , depend de la constitution de chacun des deux composants de ce 
couple. De simples variations quantitatives apparaissent qui tiennent 
aux oscillations habituelles des influences du milieu exterieur ou intra- 
uterin; ces variations n’influent pas sensiblement sur le resultat final. 
Le role de la constitution des parents est particulierement marque dans 
certains cas de croissance un peu anormale et ou l’on voit les memes faits 
























PRODUITS SUCCESSES D’UN MEME COUPLE 


305 


se reproduire dans les portees successives. Ainsi, le couple 31 donne des 
produits a croissance extremement rapide: 



1 tRE PORTAE 

2 feME PORTAE 

Petits 

Grammes 

Petits 

Grammes 

1 ere pes6e. 

1 

2.4 

3 

1.83 

2 eme pes6e. 

1 

6.20 

3 

5.66 

3 eme pes6e. 

1 

8.25 

3 

9.25 

4 £me pesee. 

1 

11.75 

3 

13.25 

5 eme pes£e. 

1 

18.00 

3 

15.21 


En 5 semaines, ces souris atteignent un poids que le plus grand nombre 
des souris n’atteignent qu’en 8 ou 10 semaines. 

La conclusion se degage done d’elle-meme. 11 ne s’agit pas d’examiner 
l’ordre de primogeniture des descendants d’un couple, mais l’etat des 
composants de ce couple et les conditions d’elevage. Celles-ci pourront 
certainement modifier celui-la; mais la question est alors toute differente. 





















THE IDEAL FAMILY HISTORY 


HOWARD J. BANKER 

Eugenics Record Office, Cold Spring Harbor, New York 

The preparation of a family history while possessing many of the elements 
of historical method differs so greatly from all other forms of historical 
writing that it would seem to be entitled to a special designation of its 
own. The term Genealogy is correctly used in too restricted a sense so 
that the authors of family histories are constantly constrained to use 
qualifying terms in the titles of their works. On the analogy of the term 
biography we would suggest ecography as an appropriate expression and 
sufficiently broad in its implications. 

Family history is much more closely allied to biography in every way than 
it is to history. In fact it is essentially a series of related biographies bound 
together by a genealogical net with only an historical background. Since 
this genealogical net is inherent in the family and is predetermined by 
biological laws which we can not change it constitutes a framework 
or skeleton that gives fundamental form to the family history. It forces 
certain very positive restrictions upon the writer of the family history and 
unless he has made himself master of the genealogy he is almost sure to be 
thrown if he attempts to ride this Pegasus. As most family historians are 
untrained, inexperienced and unacquainted with the best models of eco- 
graphic writing we find our genealogical libraries filled with a mass of poorly 
organized and undigested material from which it is sometimes possible to 
pick out fragments of more or less value. 

But it does not appear to us that even trained and experienced profes¬ 
sionals have always clearly comprehended the true nature of a family 
history study so as to treat it in a form that will give it its greatest value. 
The chief difficulty arises from the vague and ill defined use of the term 
family. In the broadest sense a complete family history never has and 
never will be written. The task is impossible. The material is not avail¬ 
able and if it were no life would be sufficiently long to compass the work, 
to say nothing of the historian becoming lost in the endless ramifications of 
the family network. The fundamental problem of the historian then is to 
set the limits or the scope of his treatment so that it will include a natural 

306 


THE IDEAL FAMILY HISTORY 


307 


and unified family group. This apparently simple and obvious fact is, 
however, rarely attained even in the most elaborate and expensive family 
histories. Nearly all are not only fragments but they are distorted and 
often sadly mutilated fragments. 

Two considerations are apt to control in determining the scope of a 
family history study neither of which is of the highest importance. The 
first is to trace back to the earliest possible ancestor and the second is to 
restrict the work to those lines that afford the greatest mechanical facility 
for the research. In the prosecution of the first object much labor and some¬ 
times heavy expense is involved including the final publication of scores of 
pages of reproductions of old documents whose only value is that they 
furnish more or less certain evidence that leads to the conclusion that 
Thomas Tracy, the emigrant, was in all probability the son of William 
Tracy. The mountain has labored and brought forth a mouse. From any 
point of view of what great value is it to know that the father of Thomas 
was called William? Do not misunderstand me. I am too much of a gene¬ 
alogist not to appreciate the satisfaction one may feel in putting his finger 
squarely on the name of an ancestor one generation farther back. The 
question is, in satisfying this instinct have we not sacrificed other facts of 
as great interest and perhaps of even greater value? It is very well known 
that historical writings often suffer from distorted perspectives and no form 
of historical writing is likely to suffer more seriously from such distortion 
than is the family history. 

Having run to cover the remotest ancestor the historian commonly 
proceeds to review more or less completely the descendants of the common 
ancestor and the emphasis here is frequently on less. It is very common 
practice to restrict this part of the family history to the descendants by the 
male line, a purely mechanical distinction but serving to facilitate research 
by confining the labor to the tracing out of a single surname. Every 
Berlove is grist for my mill. It is to be admitted that this greatly reduces 
the labor and expense of research as well as the size of the published 
volume. However, if we be not blinded by a false genealogical pride, this 
question must persistently intrude itself: Is the discovery of the name of 
the father of Thomas Tracy and the publication of all the documentary 
evidence therefor of greater interest, value or importance than the recording 
of the historical narrative of all his descendants including the female as well 
as the male lines? If both could not be included in the family history have 
we not sacrificed the greater for the less? Is not such a family history not 
only a fragment but a mutilated fragment for the line of limitation cuts 
continually right through the most intimate and vital relationships? 


308 


HOWARD J. BANKER 


Such works may perhaps be called family annals but they lack the es¬ 
sential elements of a family history. They have not only a distorted 
perspective but they are seriously lacking in the fundamental requirement 
of all historical writing, unity. Even those family histories which include 
all lines of descent both male and female are still universally lacking in a 
highly important element of unity. The consorts of the so called “ blood 
line” are commonly introduced with scarcely a comment as to their origin. 
Whether we view the subject historically, socially or biologically it must be 
recognized that the consort is just as vital and plays just as significant a 
part in the family history as does the old stock parent. They twain become 
one flesh. Both are essential to the unity of the home. Moreover the 
later descendants for whom presumably the history is written are just as 
greatly interested in the one parent as in the other and will not get a right 
conception of the influences which molded their family history unless an 
account is given of the consort’s connections. The old and wide spread 
conception of a family tree belongs to the thought of an obsolete age. 
In conformity with social as well as scientific ideas of the present the family 
group must be conceived as a more or less intricate network. 

Therefore, to secure, fundamental unity, to give to the family history 
the widest range of interest, and to preserve those features in the family 
network which have the greatest intrinsic value the following plan of 
structure should be more or less closely adhered to. Select a pair of con¬ 
sorts which, after a preliminary survey, seem most suitable for the purpose 
as planned. This may be determined by their prominence but is likely to 
serve the purpose better if chosen because of their central location within 
the family net which it is intended the history shall cover. Trace out their 
ancestry and ancestral fraternities in all lines for at least three generations. 
Then take up the descendants in the usual way following the order of primo¬ 
geniture. With each consort, however, include the ancestry and ancestral 
fraternities for at least three generations. The historian should not be 
satisfied with less than this if the material can be obtained. Whether he 
should extend his work farther he must be his own judge, the fundamental 
principle being always to work centrifugally from a selected center including 
so much of the families of consorts as to give to the whole work a family 
unity. By this method of treatment the living members of the family will 
have a much larger interest in the work as a whole, the family history will 
be more unified and brought into better perspective, the social and other 
relations of family contemporaries can be more clearly brought out and the 
distribution of hereditary characteristics will be more strikingly and 
correctly displayed. 


THE IDEAL FAMILY HISTORY 


309 


In such an arrangement of the family history it is evident that we have a 
central family stock nearly or quite complete surrounded on all sides by 
those familial factors and influences which have most vitally affected it. 
We have intimated that in this boundary region the historian may extend 
his limits according to his own judgment. It must be recognized, however, 
that too great an extension of the history into this outer region will defeat 
its own ends. First, the work will soon become unwieldy as well as un¬ 
interesting. Second, there is likely to be a loss of perspective when the 
border of the design becomes larger than the central figure. Third, the 
unity of the plan fades out in the unending diffuseness. We may soon reach 
the point where the central conception is dissipated and we are no longer 
dealing with a family but with a race. 

The above comments apply just as distinctly to overexpansion vertically 
as laterally. In the downward direction we are stopped by the impassable 
boundary of the generation of the unborn, but there is no definitely fixed 
limit to the ascending series and there are peculiar fascinations in this 
direction. A little consideration will show, however, that there are limits 
beyond which it is of no practical value to go. If any of us could trace out 
his ancestors for ten generations he would find that he had over a thousand 
and the next generation alone would add a thousand more. From our 
present understanding of the biological mechanism for the transmission of 
hereditary traits it seems practically impossible that one should have 
inherited from all of these last thousand ancestors. In other words, it is 
all but certain that many of these ancestors of only three to four hundred 
years ago are no more related to some of us, their so-called descendants, 
than if they had been Fiji Islanders. Moreover, as to their having any 
influence on our lives through the family life or by means of their ideas or 
the environment which they established at the family hearth, it seems 
clearly evident that it can be no greater at this distance than the influence 
which we receive from any of their immediate contemporaries to whom we 
can trace no genetic relationship. In fact, when we trace our ancestry 
back ten generations or more we find ourselves dealing with a population 
and not with specific ancestors. Eventually, I am the descendant of a race 
and not of a person. # It does not seem, therefore, to be of any vital sig¬ 
nificance as a rule to extend a family history over a period of more than ten 
generations. If there is significance in a greater series it is a special case 
due to the operation of some selective agency and becomes of interest as do 
all exceptions. 

Thus far we have considered only the principles determining the plan 
or framework of the ideal family history. The next step is to give the plan 


310 


HOWARD J. BANKER 


a concrete form adapted to the particular family to be studied. This is the 
strictly genealogical part of the work, but is essential. It consists of an 
enumeration of the members of the family with reference to their mutual 
relationships, each individual being identified by his name, date of birth, 
marriage and death and his place in the pedigree. All such data are simply 
means of identification and have no other significance. The relationships 
can be expressed in some cases by a numerical sequence or a system of 
tabulation, but by far the most satisfactory method where the family 
network is large and more or less complex is the pedigree chart. One might 
as well attempt to study the intricacies of a military campaign without a 
map as to study a family history without a chart. 

Having settled on the plan or framework of the family history, the next 
problem is to put flesh on these dry bones that they may live. For this 
purpose each individual as far as possible must be clothed with personality. 
We are any of us interested to know that John Howland or Martha Spencer 
was one of our ancestors but how much greater is our interest if we can have 
pictured the appearance of John or Martha, can know more distinctly how 
they conducted themselves and in what they had their interests. After all, 
these are the things we really wish to know and these are the things that have 
scientific as well as historical value and personal interest. In the matter, 
however, of trying to reproduce this living personality the average family 
historian is too persistently looking backward. He has too much of the 
instincts of the antiquarian. The remotest and least known ancestor 
usually receives the largest attention, while those members of the family 
with whom the historian is well acquainted or concerning whom informa¬ 
tion may be readily acquired are passed over with scarcely more than the 
mention of the name, because forsooth they have not an antique flavor. 
A family historian does the greatest service to his family when he writes 
fully of those matters of which he knows the most. With face turned 
forward he should gather and record the information which he alone is in a 
position to preserve that the generations to come may have accurate por¬ 
traits of their kin who passed away before their time. 

Occasionally a father or mother does this for the children and these 
accounts though brief and often limited in scope are usually the most valu¬ 
able of family chronicles. Such family narratives also approach more or 
less crudely the centrifugal type of family history which this paper has 
endeavored to present. The narrator is commonly the center and passes 
entirely around the circle of his family connection, describing each one and 
telling of their interests, achievements, personal appearance and pecu¬ 
liarities; their interrelations and reactions to each other and to their en- 


THE IDEAL FAMILY HISTORY 


311 


vironment. Now if we can conceive of a score or more of such raconteurs 
properly grouped and united by the bonds of a common family network we 
have the finished structure, the body of the ideal family history. But not 
yet is it a living body. The highest type of family history still awaits the 
genius to give it full expression,—the man with comprehensive under¬ 
standing, with broad conceptions and with profound philosophical insight, 
who is able to grasp this family group as a living entity, can analyze its 
complex internal reactions, the interplay of heredity and personality and 
then project it as a vital element in its social and physical environment, as a 
potential factor in the historical development of the race. This is the goal 
toward which as genealogists and ecographers we should strive. This is 
the ideal which should more largely animate and inspire our genealogical 
and patriotic societies. Preserve the past with all diligence but remember 
that tomorrow, today will be yesterday. 


THE CONIFICATION OF SOCIAL GROUPS: EVIDENCE FROM 
NEW ENGLAND FAMILIES 

FREDERICK ADAMS WOODS 
Massachusetts Institute of Technology 

The word conification, which appears in the title of this paper, has, as 
far as I know, never been used before. The introduction of a new word into 
a language requires an explanation, if not an apology. If any apology is 
needed, my hope is that it will be accepted rather in the nature of an excuse, 
because no other word could be found that at all well expresses the process 
to be described. The word “ conif ormation” is probably somewhat clearer, 
and is to be found in the Century Dictionary, though not often used. It 
means the formation of a cone. But it does not well express a general proc¬ 
ess. Just as we might speak of the solidification of groups and that groups 
solidify, so it is necessary to say that social groups at times show conifica¬ 
tion or conify, i.e., tend to point upward, like a cone. 

If all the persons of a social group come together closely, in a mental or 
spiritual way, as for instance in a time of religious revival, or during a period 
of war, the group may be said to solidify. If a different process is to be 
described and some people, figuratively speaking, climb on the shoulders of 
others, the individuals form what may well be represented by the word 
conification. 

In the accompanying figures we see in profile (or cross section) first a 
cone of flattened proportions like a fiat turnip (fig. 1). This graph repre¬ 
sents a group in which the individuals are, in any attribute under discussion, 
very much alike. Each individual might be represented by a single particle 
or dot. A few are near the top of the pile, a few are near the bottom, while 
the vast majority find their places in the broad median zone which consti¬ 
tutes the greater part of the structure. This median zone is emphasized 
by the lighter band. 

The next diagram (fig. 2) shows a change. Some have succeeded in 
getting themselves into a higher position and some have been forced into 
a position even lower than before in relation to the mass. The third figure 
represents the same process carried to a greater degree. 

Such graphs may be used to illustrate various special aspects of group 
differentiation, such as wealth, natural intelligence, outward achievement, 

312 


CONEFICATION OF SOCIAL GROUPS 


313 


physical strength, stature, in fact, any trait that can be measured. These 
cones can also be used to illustrate certain unmeasurable or vaguely measur¬ 
able social and national differences. For instance, if one country is con¬ 
sidered more aristocratic than another, the fact can be represented graphi¬ 
cally by two cones, one of which is more pointed at the top than the other. 
Here, of course, the exact forms can not be plotted, but the idea can be put 



FIG. 1 FIG. a 



t 


FIG. 3 

into the form of a drawing,—a cross section of an object of three dimensions, 
—a solid object familiar to the eye and easily reproduced in the imagination. 
Conification is the transformation from the form of a flat turnip to one of 
the pointed variety. Nations can then be thought of, not as being composed 
of people who are all alike (which certainly they are not), but as composed 
of differentiated interacting particles. Thus, France and Germany before 



314 


FREDERICK ADAMS WOODS 


the war might be represented by figure 2 and figure 3 respectively, or 
by one and two or by one and three. Figure 1 is the most democratic, 
figure 3 is the most aristocratic. The fact that there are such differences 
and that these need to be reckoned with in discussions relating to histori¬ 
cal change is the point I wish to bring out. 

There are always or nearly always forces at work acting towards coni- 
fication or sharpening and raising the point of the social structure and there 
are always some forces acting in the opposite direction tending towards 
leveling the group. At times and in some countries these leveling forces 
have acted very suddenly, as during revolutions or conquests. The other 
or aristogenetic forces have scarcely been recognized though they are proba¬ 
bly always active in civilized countries during normal periods of growth. 
Their activities are to be expected as corollaries of certain scientifically 
recognized facts. One of these facts is that, by and large, through the liv¬ 
ing world, like tends to mate with like, and this is true in a general way 
among civilized human beings. The children of the well-to-do and suc¬ 
cessful tend to marry within their own circle, while the poor, as a class, are 
forced to mate with the poor, and the middle classes on the average marry 
among themselves. 

The second fact is that like tends to beget like. These two forces acting 
together lead towards diversification of the population as a whole. No 
one should doubt the reality of these forces. The only question is, do these 
inner forces, assortive mating combined with heredity, outweigh the en¬ 
vironmental, leveling and democratic forces and produce a measurable re¬ 
sult? Do nations and social groups, if left to themselves, actually conify or 
in other words become more aristocratic? There is evidence that they do. 

First, there is evidence from the study of royal families. These fami¬ 
lies were originally formed through a process of selection, and a survival 
of the fittest in two important attributes, namely, success in war and 
government. 

The early kings of the Anglo-Teutonic tribes were either elected for merit 
or made themselves kings by force of will and ability. Their children in¬ 
termarried and their descendants for hundreds of years did produce on the 
average a very high proportion of men naturally gifted in the arts of war and 
government. There are about twenty persons of unquestioned genius, 
for the most part closely related, in a total population of less than one 
thousand. 1 

1 Frederick Adams Woods, Heredity in Royalty, 1906, and The Influence of Monarchs, 
1913. 


CONIFICATION OF SOCIAL GROUPS 


315 


Second, there is evidence from the British Dictionary of National Biog¬ 
raphy. It appears that, from the earliest times to the beginning of the 
nineteenth century, the contribution to eminent achievement made by 
the sons of craftsmen, artisans and unskilled laborers yielded 11.7 per cent 
of the total number of names. The same class born during the first quarter 
of the nineteenth century only produced 7.2 per cent of the total and I 
have found that the second supplement which contains names of persons 
born chiefly during the second quarter of the nineteenth century shows only 
4.2 per cent born from the same working classes. Thus the great mass of 
the population appears to be more separated off from the upper classes than 
formerly, at least in this one particular respect, namely the power of pro¬ 
ducing men of exceptional talent. 2 

Third, a study of the history of representative New England families 
shows this conification actually taking place. 

There is perhaps no small geographical territory in the world where local 
family records can be so easily and completely obtained as in Boston and 
vicinity. Most of the manuscripts have been printed and are to be found 
in the library of the New England Historic Genealogical Society. Wyman s 
history of Charlestown, Bond’s Watertown, Paige’s Cambridge, and Butler’s 
Groton (the last supplemented by the numerous monographs of the late 
Dr. Samuel A. Green) carry local history almost to the final word. The 
records of births, baptisms, marriages, deaths, wills, deeds, town meetings, 
and military enlistments have been so thoroughly gone over for the earlier 
generations, that practically every person is known. The marriages are 
nearly all known, and the social status of the individuals and that of their 
parents are also known. Unfortunately we possess few facts concerning 
the inner traits and characteristics of most of these people, but we do know 
enough to be able to measure their comparative achievements, and their 
occupations and degrees of public service can almost always be determined. 
Private genealogies rarely contain anything bad about any one, but their 
writers are very eager to include all the good. Therefore, if there is nothing 
more to be found about a man than the dates of his birth, marriage, and 
death, we may be reasonably certain that, no matter how excellent he may 
have been in character, or even in natural ability, as far as his relation to the 
community was concerned he was not an influential man. The social order 
demands achievement, and towards merit without achievement its indiffer¬ 
ence is but natural. It is the relative achievements as well as the social 
positions of these people that can be studied with comprehensive results. 

2 See Alleyne Ireland, Democracy and the Human Equation, p. 139, for the application 
of this matter to the theory of democracy. 


316 


FREDERICK ADAMS WOODS 


Also we may follow the intermarriages between the different social classes 
and estimate fairly well the class distribution of wealth. 

As has been always recognized, the great majority of the early settlers in 
Massachusetts Bay belonged to the class, at that time alluded to as the 
yeomanry. There was a sprinkling, say 5 per cent, of gentry, and a very 
small and almost negligible number of vagrants or others now called “ un¬ 
desirables,” but there were probably no breeds of degenerates such as were 
formed in certain hill-towns in later days. There were no tribes of “ Jukes” 
or “Nams” at the start. 

The gentry was made up in part by those who belonged to the recorded 
gentle families of England,—in other words,had the right to a coat of arms. 
There were surprisingly few such families and modern critical research has 
shown that the number probably does not exceed seventy-five for all New 
England. There were others who must be counted as belonging to the 
gentry, owing to their wealth or their high governmental offices or their 
membership in the then dominating theological class. College graduates 
and members of the learned professions may also be included in the 
class called gentry. Artisans and small shopkeepers may be classed with 
yeomen. 

Between the gentry and ordinary yeomanry there existed certain indi¬ 
viduals who, though comparatively uneducated, according to our present 
standards, were nevertheless real and practical leaders, and doubtless in¬ 
telligent, enterprising, and courageous. They were the typical officers of 
the militia and pioneer founders of new towns on the frontier. These, if 
yeomen, I have called “ officer yeomen,” and we can often even speak of 
whole families as “ officer yeoman,” when a considerable number were 
closely related. The Massachusetts Society of Colonial Dames has listed 
in its Register a very large proportion of such persons, as well as those of 
greater prominence. These are considered as “eligible” ancestors for the 
election of their descendants in the Society. 

Starting with this Register I have taken out the names of all families 
settled prior to 1692 who had three or more members now eligible as ances¬ 
tors for Colonial Dames and at the same time lived in Boston or Charles¬ 
town, Watertown, Roxbury, Cambridge, Concord, Woburn, or Groton. 
This furnishes a list of seventy-one families or male-lines. Out of this 
seventy-one, forty (whose names are here given in the footnote 3 ) were chosen, 

3 Families of the gentry and “officer yeomanry” in Boston and vicinity whose pedigrees 
have been studied and included in this research: 

Adams, 23 Appleton , 2 ’ 3 Bigelow, 3 Boylston, 2 Bradstreet , 2 Breck, 2 Brooks, 3 Bulkeley , 
Chandler, 2 Chauncy, 2 Cheever, Converse, Coolidge, 3 Dudley, Farwell, French of Cambridge, 


CONIFICATION OF SOCIAL GROUPS 


317 


partly because of personal interest and knowledge that these families were 
today represented by descendants of more or less prominence, and partly 
because they happened to be connected with those taken at the start, so 
that the inclusion of one led to the inclusion of another, as the pedigree 
charts expanded, until a sufficiently great group of names had been obtained. 
This has now exceeded 3000 and it does not seem necessary to carry it further. 
There is no reason to suppose that the remaining thirty-one families whose 
names are also given in the second footnote, 4 and whose records were not 
included merely from lack of sufficient time, would in any way affect the 
present conclusions. 

It has been the aim in this research to select by an objective method a 
characteristic leading class in America in the early days to compare with a 
similar class in the nineteenth and twentieth centuries. The great strik¬ 
ing fact is that the upper portion of the body-social, as represented by 
families of the gentry, is more separated from the mass today than it was in 
colonial times. At least in two very important matters it is so separated. 
The first is wealth and the second is intermarriage. These both mean 
conification. 

As regards the distribution of wealth among the four classes considered, 
we know that the total proportion of paupers and dependents was very 
small in the early days. The average yeoman possessed, as shown by in¬ 
ventories of wills, about £100 to £300. If a man had more than £300 
personal property he might, with propriety, wear lace. At least there was 
at one time a law passed forbidding the use of lace to persons not possess¬ 
ing £300 personal property. Well-to-do yeomen, officer-yeomen, and gen¬ 
tlemen in the then technical sense of the word often possessed from £500 
to several thousand pounds. Rev. Peter Bulkeley, the founder of Concord, 
a member of a very old English family and one of the most wealthy men in 
the colony, brought with him from England £6000. Robert Kayne of 

etc., Frothingham, 3 Hancock, 2 Hunnewell, 3 Johnson of Woburn, Lawrence of Groton, 3 
Learned, Lynde, Mather, Minot, 2,3 Oliver , 2 Phillips, 2 ' 3 Prescott, 2 Quincy, 2 ’ 3 Russell , 2 
Ruggles, Saltonstall, 2 ’ 3 Sparhawk, 2 Stearns of Watertown, Symmes, Tarbell, Willard, 
Winthrop, 2 Woods of Groton, Wyman. The thirteen names selected as showing special 
prominence in the first century are in italics. Those marked 2 were selected for the period 
1721-1820. Those marked 3 for the period 1820-1921. The middle period, 1721-1820, 
has not been finished. A rough estimate indicates that the change from the earlier 
century was not great, until about the beginning of the nineteenth century. 

4 The following families have not as yet been studied: Allen, Barrett, Bowers, Clapp, 
Clark, Cotton, Craft, Danforth, Davenport, Denison, Eliot, Fiske, Foster, Gookin, 
Hutchinson, Leverett, Mason of Watertown, Pynchon, Sargeant, Savage, Sawtell, Sewall, 
Shrimpton, Sprague, Stoddard, Stone, Tyng, Thurston, Weld, Wheelwright, Whittingham. 


318 


FREDERICK ADAMS WOODS 


Boston, one of the richest men in the first generation of settlers, left an 
estate in 1656 valued at £2569. Captain Thomas Brattle died in 1683, 
leaving an estate of £7827, which according to Savage was, “ probably the 
largest in New England.” Thus we see that the richest men were not more 
than fifty times as rich as the average. By the middle of the eighteenth 
century the differences had become somewhat greater. In Dwight’s 
Travels it is said that Col. Elisha Doane of Wellfleet and Boston, and 
Thomas Boylston, of Boston, were estimated by their fellow citizens the 
two richest men in the province of Massachusetts Bay. Doane’s fortune 
was estimated at £100,000. This was perhaps 300 times that of the average 
citizen. 

In a gossipy little book called the “Rich Men of Massachusetts,” pub¬ 
lished in 1851, we find a number of persons estimated as worth a million 
dollars or more. The richest man is placed at three millions. This must 
be more than 600 times the average. The differences increase as we ap¬ 
proach the present day, during which time many persons in New England 
have left estates valued at twenty millions or more, or several thousand 
times the average. If we consider the United States as a whole, the very 
richest men today, those who are worth $100,000,000 or more, are certainly 
as much as 10,000 times to 100,000 times as rich as the average. There can 
be no question that, as far as the distribution of wealth is concerned, there 
has been in America a process of conification. The average has risen some¬ 
what, but the point of the cone has risen faster than the mass. If the mass 
be represented in the graph, as six inches high, the top of the drawing 
would have to be carried up at least 5000 feet in the air. 

Now let us examine the question of social differentiation as shown by the 
intermarriages of the different classes. We are apt to think of the colonists 
of the seventeenth and eighteenth centuries as very particular about their 
social rank. In outward forms they may have been. They were very 
particular about precedence in seating in church and somewhat so in ar¬ 
ranging names in college catalogues, though not rigidly so, as a study of the 
early Harvard class lists will show. But when it comes to the matter of 
marriage and intermarriage between the three different social classes, we 
find a surprising lack of just this class distinction. There is certainly no 
better test than intermarriage between classes to show whether different 
castes mingle freely in a social way. At least this is the biological or eugenic 
test par excellence, as it is the one which determines the heredity of the next 
generation. 

It is true that even in the earliest period there were certain families that 
may be spoken of as belonging to the gentry. These naturally intermarried 


CONIFICATION OF SOCIAL GROUPS 


319 


to a considerable extent, just as is the custom everywhere. Characteristic 
families of this description are represented by those of the governors, Dudley, 
Bradstreet, Winthrop, and Saltonstall, the wealthy Russell family of 
Charlestown, and the ancient pedigreed and theocratic families of Bulkeley 
and Chauncy. There are included in this study thirteen such preeminent 
families (prior to 1721) out of the total of forty. But in scarcely one of 
these is there not an instance of intermarriage with yeomanry, and usually 
many such examples can be counted. 

At this point I wish to express my indebtedness to the well-known geneal¬ 
ogist, Mr. J. Gardner Bartlett, who has gone over most of the data con¬ 
cerning the early settlers and has assisted in classifying the persons into the 
three grades here considered. As Mr. Bartlett has written a large number 
of histories of New England families, containing thousands of names, and 
has also specialized in the English ancestry of the early settlers, he is pecul¬ 
iarly fitted to aid in this way. My own feeling is that no one else could have 
done this work better than he, and I have used Mr. Bartlett’s classification 
rather than my own whenever there has been a question. Thus a point is 
gained towards objectivity and impartiality, always a desirable feature in 
any historiometric work. Other things equal, it is better to report on other 
people’s measurements than on one’s own, especially when personal theory 
enters into the matter. 

In this case I did have the theory, outlined in a chapter published in 1913, 
in the “Influence of Monarchs,” that social conification, must in civilized 
societies be a normal process, but this is the first local evidence obtained. 
Here we really see the forces of aristogenesis sprouting up against and 
through the forces of democracy, making themselves obvious and easily 
susceptible of statistical proof. 

Out of the forty families (male-lines) studied, thirteen have been selected 
as undoubtedly belonging in the highest social position during the period 
prior to 1721. These are italicized in the footnote on page 316 already 
referred to. 

A detailed analysis cannot be given here, but in a word, the result is that 
out of 204 cases of marriage, in this early period, 48 or 23.5 per cent were 
marriages of the children of the gentry with the children of yeomen or, in 
other words, cross-marriages between the upper and the lower of the three 
social grades here considered. 

After finishing this bit of statistical inquiry, I found, on re-examining the 
“ Memorial History of Boston” (a standard work in four volumes prepared 
by a group of scholars and specialists), that a list of forty families socially 
prominent in the seventeenth century had been prepared by W. H. Whit¬ 
more, author of the chapter dealing with the social life in Boston. 


320 


FREDERICK ADAMS WOODS 


Here were a good many names not included in my research. These 
were families strictly associated with Boston, and not with the outlying 
towns. Fearing that my own list did not represent a true selection of the 
most important names, and so might falsely overestimate the amount of 
cross-marriage of gentry with yeomanry, I looked up the records of the 
first seventeen of these families and found that the first hundred marriages 
yielded again approximately 23 per cent of cross-marriages between the 
upper and the lower of the same three grades. 

Let us now contrast these conditions with those during the last hundred 
years. Twelve families are again selected from the same forty whose pedi¬ 
grees have been traced. These are the families who appear to have had dur¬ 
ing the last century (1821-1921), the greatest amount of social and financial 
prominence. They are marked 3 in the first footnote. A few others might 
equally well have been selected from my list of forty pedigrees, but they 
have been omitted because their members have to a great extent moved 
away from Massachusetts or have now only a few representatives in the 
male-lines. Of course dozens of families, if selected for present day promi¬ 
nence, might be added, but that would mean another research. 

The result of looking up the records of marriage and parentage in the 
twelve families that happen to be included shows within the last century 
that out of 152 marriages at least 143 or over 94 per cent, are certainly 
within their own class. There are not more than two instances of known 
marriages distinctly outside the class to which these people belong. The 
parentages of a few of the persons who happen to fall within this group are 
very difficult to trace in any printed records; probably most of them came 
from a class between the two extremes, such as, for the colonial period, we 
have called officer yeomen. But it is not necessary to know the ancestry 
of these few persons. Even if they are considered as Y (or similar to yeo¬ 
manry of early period), the percentage of cross-marriages gentry with yeo¬ 
manry has been reduced from 23 to less than 6. If we look at it another way 
out of 204 marriages from families of the gentry in the earliest period, 119 
or only 58.3 per cent were within their own grade. During the third period, 
subsequent to 1820, at least 94 per cent have been strictly within the same 
class. The tendency towards casts marriages has increased markedly, and 
has increased recently. 

These facts taken in conjunction with the increasingly uneven distribu¬ 
tion of wealth prove that social conification does take place. It is probably 
inevitable whenever a population, at first composed of comparatively similar 
persons, lives in a territory where inheritable wealth can be acquired. 
It is probably working all the time in all civilized countries, though it may, 


CONIFICATION OF SOCIAL GROUPS 


321 


since it requires several generations to show its results, be masked super¬ 
ficially or buried under the wreckage of revolutionary debacle. 

Recently in Germany, Austria, and Russia, for instance, there have been 
gigantic examples of the breaking down of cones. Probably many social 
cones tend to break, some for one reason, some for another, as they become 
overconified, or too much pointed at the top. But they certainly tend to 
form, and it is merely to bring forth statistical and historical evidence of 
such formation that the present paper has been prepared. 

The process of conification is hastened and intensified from the correla¬ 
tion that exists between social status and intellectual achievement. This 
correlation has been demonstrated in royalty, in nobility, in American 
families connected with persons in the Hall of Fame and in other material. 
The present research was not undertaken to prove the existence of such a 
correlation, since this had been known and recognized, but it is not without 
interest to see how many persons show high intellectual attainments in a 
group of families selected solely for wealth and social position since 1821. 

This group of less than two hundred persons yields the following names at 
or above the standard of inclusion of Who’s Who in America: Charles 
Francis Adams born 1809. Charles Francis Adams born 1835, Henry 
Adams born 1838. Brooks Adams, Dr. Henry J. Bigelow, Phillips Brooks; 
Algernon, T. Jefferson, J. Randolph, Julian L. and Archibald C. Coolidge; 
Rev. Nathaniel, Rev. Octavius B., Rev. Paul and Louis A. Frothingham; 
Amos A. (born 1814) and Bishop William Lawrence, Dr. Charles S. Minot, 
Wendell Phillips. 

When one considers that not one adult male in a thousand attains any 
such distinction and that here out of a group of less than two hundred men 
and women, one can easily count nineteen, one sees how undoubtedly 
positive this correlation is. 

The conclusion appears unescapable that no matter how much we may 
contemplate environmental forces, making for equality and democracy, 
here the real result has been in the opposite direction, namely, class differ¬ 
entiation and conification, due presumably to the accumulating forces of 
assortive mating, heredity, and the possibilities of transmitted wealth. 

GENTRY, FIRST PERIOD (PRIOR TO 1721) 

Appleton 

Gentry X Gentry 

m dau. Rev. Jesse Glover 
m dau. William Paine of Ipswich 
m (2) dau. John Oliver of Newbury' 


Capt. John b. 1622 

Col. Samuel b. 1625 
« u 


322 


FREDERICK ADAMS WOODS 


Sarah 

m 

Rev. Samuel Philips 

Judith b. 1634 

m 

Samuel Rogers of Ipswich 

Judith 

m 

Samuel Wolcott of Wethersfield 

Samuel b. 1654 

m 

Elizabeth Whitingham 

Oliver b. 1672 

m 

dau. Tobijah Perkins of Topsfield 


Gentry X Officer Yeoman 

Major Isaac b. 1664 

m 

dau. Thomas Baker of Topsfield 


Gentry X Yeoman 

Martha b. 1620 

m 

Richard Jacobs of Ipswich 

John b. 1660 

m 

(2) Elizabeth Dutch of Ipswich 

Joannah 

m 

Mathew Whipple 

Bradstreet 


Gentry X Gentry 

Hon. Simon b. 1663 

m 

(20, widow Ann Gardner) 

u ft 

m 

dau. Gov. Thomas Dudley 

Dr. Samuel H. C. 1653* 

m 

Mercy Tyng 

Dorothy 

m 

Rev. Seaborn Cotton 

Sarah 

m 

Richard Hubbard, H. C. 1653 

Rev. Simon 

m 

dau. Rev. John Woodridge 

Dudley 

m 

Ann widow of Theodore Price 

John 1653 

m 

dau. Rev. William Perkins 

Mercy b. cir. 1667 

m 

Dr. James Oliver 

Lucy 

m 

Hon. Jonathan Remington 

Dudley b. 1678 

m 

Mary Wainwright 

Simon b. 1682 

m 

dau. Rev. Joseph Capen 


Gentry X Officer Yeoman 

Hannah 

m 

Andrew Wiggin of Exeter, N. H 

Mercy 

m 

Maj. Nathaniel Wade 

Rev. Simon 

m 

Mary Long dau. John of Charlestown 

Sarah 

m 

(2) Maj. Samuel Ward 


Bulkeley 
Gentry X Gentry 

dau. Rev. John Jones 
Rev. Joseph Emerson 
(2) John Brown Esq. 
Capt. Ephraim Flint 
Mrs. Rebecca Minot 
(2) Silence Jeffrey 
dau. President Chauncy 
Rebecca Talcott 


Thomas b. 1617 

m 

Elizabeth 

m 

u 

m 

Jane 

m 

Capt. Joseph b. 1670 

m 

« u 

m 

Rev. Gershom 

m 

Peter 

m 


* H. C. = Harvard College graduate. 




CONIFICATION OF SOCIAL GROUPS 


323 


Hon. Peter 
Rebecca 
Dorothy 
Edward 
Rev. John 


Rebecca b. cir. 1695 
Catherine b. cir. 1660 


Gentry X Officer Yeoman 

m. dau. Lt. Joseph Wheeler 
m Jonathan Prescott Jr. 
m Lt. Thomas Treat 
m Dorothy Prescott dau. of Jonathan 
m Patience Prentice 

Gentry X Yeoman 

m Joseph Hubbard 
m Richard Treat 


Sarah 

Katherine 

Abigail 

Sarah 

Charles d. 1711 


Abigail 

Rev. Nathaniel 


Rev. Nathaniel b. 1639 


Chauncey 

Gentry X Gentry 

m Gershom Bulkeley 
m Rev. Daniel Brewer 
m Dr. Hudson 
m Rev. Samuel Whittelsey 
m dau. Judge John Walley 

Gentry X Officer Yeoman 

m Edward Burroughs 
m Sarah Judson 

Gentry X Yeoman 

m Abigail Strong 


Dudley 


Gentry X Gentry 


Gov. Thomas 

m 

Mrs. Catherine Hackburn 

Rev. Samuel b. 1610 

m 

dau. Gov. John Winthrop 

u u 

m 

(2) Mary Byley 

Anne 

m 

Gov. Simon Bradstreet 

Patience 

m 

Maj. Gen. Daniel Denison 

Sarah 

m 

Maj. Benjamin Keaine 

Mercy 

m 

Rev. John Woodbridge 

Deborah 

m 

Jonathan Wade 

Gov. Joseph 

m 

dau. Judge Tyng 

Paul “merchant” 

m 

dau. Leverett 

Thomas “sea captain” 

m 

Abigail Gilman 

Judge Paul F.R.S. 

m 

Lucy Wainwright 

Rebecca 

m 

Samuel Sewall 

Mary 

m 

1675 Dr. Thomas Hardy 

Hon. William b. 1686 

m 

dau. Judge Davenport 

Catherine 

m 

Lt. Gov. Dummer 

Mary 

m 

Francis Wainwright 


324 


FREDERICK ADAMS WOODS 


Gentry X Officer Yeoman 


Anne b. 1684 

m 

(2) Jeremiah Miller 

Elizabeth 

m 

Hon. Kinsley Hall 

Stephen 

m 

dau. Hon. John Gilman 


Gentry X Yeoman 

Anne 

m 

Edward Hilton 

Sarah dau. Gov. Thomas 

m 

(2) Thomas Pacy 

James b. 1663 

m 

Elizabeth Leavitt 

Abigail dau. Rev. Samuel 

m 

Jonathan Watson 

Dorothy 

m 

Moses Leavitt 

Rebecca 

m 

Francis Lytard 

Samuel 

m 

dau. Jonathan Thyng 

Rev. Samuel 

m 

(3) Elizabeth 

Beyley 

m 

dau. Moses Gilman 

Timothy 

Mather 

Gentry X Gentry 

m dau. Maj. Gen. Humphrey Atherton 

Rev. Eleazar 

m 

dau. Rev. John Warham 

Rev. Increase 

m 

dau. Rev. John Cotton 

t ( « 

m 

(2) widow Rev. John Cotton 

Rev. Samuel b. 1650 

m 

dau. Gov. Robert Treat 

Rev. Cotton F.R.S. 

m 

dau. Col. John Phillips 

{( u 

m 

(2) dau. Dr. John Clark 

U it 

m 

(3) dau. Rev. Samuel Lee 

Elizabeth 

m 

Josias Byles 

Sarah 

m 

Rev. Nehemiah Walter 

Abigail 

m 

(2) Rev. John White 

Hannah 

m 

Peter Oliver 

Jerusha 

m 

Peter Oliver 

Abigail b. 1694 

m 

Daniel Willard 

Atherton b. 1663 

m 

(1) Rebecca Stoghton 

Joseph b. 1661 

Gentry 

m 

X Officer Yeoman 

Sarah Clapp 

Richard b. 1653 

Gentry X Yeoman 

m Catherine Wise 

Catherine 

m 

1651 Ebenezer Withington 

Atherton b. 1663 

m 

(2) Mary Lamb 

Samuel b. 1683 

m 

Deborah Champion 

Capt. Timothy 1681 

m 

Sarah Noyes (?) 

Lt. Joseph 1686 

m 

Phebe 

Dr. Samuel H. C. 1698 

m 

dau. Samuel Grant 



CONIFICATION OF SOCIAL GROUPS 


325 


Minot 

Gentry X Gentry 


James b. 1628 

m 

dau. Col. Stoughton 

Capt. John b. 1626 

m 

dau. Nicholas Butler (or 0) 

Elizabeth 

m 

Rev. John Danforth 

Mehatabel 

m 

(2) Soloman Stoddard 

Stephen b. 1688 

m 

(1) dau. Col. Francis Wainwright 


Gentry X Officer Yeoman 

James H. C. 1675 

m 

dau. Capt. Timothy Wheeler 

Col. Stephen b. 1632 

m 

dau. Capt. Christopher Clark 

Col. James b. 1694 

m 

dau. Col. John Lane 

Stephen b. 1688 

m 

(2) dau. Capt. John Brown 

Elizabeth 

m 

Capt. Daniel Adams 

Stephen 

m 

Trucross Davenport 


Gentry X Yeoman 

Capt. John b. 1626 

m 

(2) Mary Biggs 

Samuel b. 1635 

m 

Hannah Howard 

John 

m 

dau. Edward Breck 

Samuel 

m 

Hannah Jones 

James b. 1659 

m 

Rebecca dau. John Jones 

Mehetabel 

m 

Thomas Cooper (prob.) 

Rebecca b. 1685 

m 

Joseph Barrett 

Lydia 

m 

Benjamin Barrett 

u 

m 

(2) Samuel Stone 

Mary 

m 

Ebenezer Wheeler 

Rev. Timothy 

m 

dau. Noah Brooks 


Oliver 


Gentry X Gentry 


Rev. John H. C. 1645 

m 

Elizabeth Newdigate 

Peter 

m 

Sarah Newdigate 

Nathaniel 

m 

dau. Thomas Brattle 

James 

m 

dau. Samuel Bradstreet 

Daniel 

m 

dau. Hon. Andrew Belchor 

Sarah 

m 

Rev. Caleb Throwbridge 

Oliver b. 1677 

m 

Sarah Perkins of Topsfield 


Gentry X Officer Yeoman 

Thomas 

m 

dau. Capt. Thomas Prentice 

Abigail 

m 

Hon. Benjamin Prescott 


326 


FREDERICK ADAMS WOODS 


John 

Thomas 

Sarah 

Elizabeth 


Rev. George 
Rev. Samuel 
Samuel b. 1657 
Elizabeth 


Rev. Samuel 
Sarah b. 1691 
Mary 
Ruth 


Elizabeth 

Zerobabel 

Jonathan 

Theophilius 

Abigail 

Sarah 


Lt. Col. Edmund 

Daniel 

Judith 

Elizabeth 

Mary 

John 

Edmund b.Jl681 
Joanna 


Ruth 

Judith 


Experience 


Gentry X Yeoman 

m dau. John Sweet 
m (2) Mary Wilson 
m John Noyes 
m Enoch Wiswall 

Phillips 

Gentry X Gentry 

m dau. Richard Sargent 
m Sarah Appleton 
m dau. Rev. John Emerson 
m Rev. Edward Payson 

Gentry X Officer Yeoman 

m dau. Capt. John White 
m William White of Haverhill, N. Y. 
m Capt. Geo. Abbott 
m Samuel White of Haverhill 

Gentry X Yeoman 

m Job Bishop of Ipswich 
m Anne White 
m Sarah Holland 
m Mary Bennet 
m James Barnard 
m Stephen Mighill 

Quincy 

Gentry X Gentry 

m (2) dau. Maj. Gen. Gookin 
m dau. Rev. Thomas Shepard 
m Rev. John Rayner, Jr. 
m Rev. D. Gookin 
m Rev. D. Baker 
m dau. Rev. John Norton 
m Dorothy dau. Rev. J. Flint 
m David son Rev. Peter Hobart 

Gentry X Officer Yeoman 

m Capt. John Hunt of Weymouth 
m John Hull 

Gentry X Yeoman 

m William Savil of Braintree 


CONIFICATION OF SOCIAL GROUPS 


327 


Russell 



Gentry X Gentry 

Hon. James b. 1640 

m Mabel Haynes 

it It tt 

m Mary Wolcott 

Rev. Daniel 

m dau. Samuel Willis 

Mehetabel 

m Rev. John Hubbard 

u 

m (2) Rev. Samuel Woodbridge 

Hon. Daniel 

m dau. Capt. Charles Chambers 


Gentry X Officer Yeoman 

Elizabeth 

m Capt. Nathaniel Graves 

Mabel 

m David Jenner 

Mary 

m Capt. John Miller 


Gentry X Yeoman 

Katherine 

m William Roswell 

Maud 

m Daniel Lawrence 


Saltonstall 


Gentry X Gentry 


Nathaniel H. C. 1659 m dau. Rev. John Ward 

Gov. Gurdon m dau. James Richards 

Elizabeth m Rev. John Dennison 

“ m (2) Rev. Roland Cotton 

Richard m dau. Capt. Simon Wainwright 

Gentry X Officer Yoeman 

Nathaniel H. C. 1695 m widow of John Frizel 

Winthrop 


Gentry X Gentry 


Elizabeth 

m 

Rev. Antipas Newman 

ii 

m 

(2) Zerubbebel Endicott 

Lucy 

m 

Major Edward Palmes of New London 

Margaret 

m 

John Corwin 

Martha 

m 

Hon. Richard Wharton 

Anne 

m 

Hon. John Richards 

Waitstill 

m 

Mary d. Hon. William Browne 

U 

m 

(2) Katherine d. Hon. William Brattle 

John b. 1681 

m 

Anne d. Jos. Dudley 

Anne b. 1686 

m 

Thomas Lechmere 

Martha b. 1630 

m 

Thomas Lyon of Stanford 

Mary b. 1612 

m 

Rev. Samuel Dudley 

Adam b. 1620 

m 

dau. Rev. J. Glover 

Deane b. 1632 

m 

dau. Rev. J. Glover 


328 

FREDERICK ADAMS WOODS 

Margaret b. 1660 

m 

Jonathan Glover 

Priscilla 1669 

m 

Eliab Adams 

Mercy 

m 

Atherton Hough 

Adam b. 1676 

m 

dau. Col. John Wainwright 

Martha 

m 

Col. John Ballantine 



Gentry X Officer Yeoman 

Adam 

m 

dau. Capt. Thomas Hawkins 

Deane b. 1632 

m 

(2) widow Capt. John Mellows 

Elizabeth 

m 

Capt. Samuel Kent 


Gentry X Yeoman 

John (Fitz John) 

m 

dau. George Tongue of New London 


THE MAYFLOWER PILGRIMS 

SARAH LOUISE KIMBALL 

What profit pedigree or long descents 1 2 
From farre-fecht blood, or painted monuments 
Of our great-grandsire’s visage? ’Tis most sad 
To trust unto the worth another had 
For keeping up our fame; which else would fall, 

If, besides birth, there be no worth at all. 

For, who counts him a gentleman whose grace 
Is all in name, but otherwise is base? 

Or who will honour him that’s honour’s shame, 

Noble in nothing but a noble name? 

It’s better to be meanly born and good, 

Than one unworthy of his noble blood: 

Though all thy walls shine with thy pedigree , 

Yet virtue only makes nobility? 

Then, that this pedigree may useful be 
Search out the virtues of your family; 

And to be worthy of your father’s name, 

Learn out the good they did, and do the same: 

For, if you bear their arms, and not their fame, 

Those ensigns of their worth will be your shame. 

We claim for the descendants of the Pilgrims an especially pure, high 
minded type of character, exalted spiritually in an eminent degree. 0 
Had it not been for the faith and courage of that little band of English 
exiles, the founders of Plymouth Colony, America would not have developed 
along the lines which have placed her in the lead among the nations of the 
world. 

Prior to the landing at Provincetown, Cape Cod, on December 21, 1620, 
and on November 21,1620, the famous Compact drawn up by William Brad¬ 
ford was signed in the cabin of the Mayflower. 

Y® COMPACT 

IN THE NAME OF GOD, AMEN 

We whose names are underwritten, the Loyall Subiects of our dread soveraigne Lord 
King lames, by the Grace of God of Great Britaine, France, and Ireland King, Defender 
of the Faith, &c. 

1 Old Latin Verses quoted in Burke’s “Patrician,” iii, 63-4. 

2 Motto of California Genealogical Society. 

329 




330 


SARAH LOUISE KIMBALL 


Having under-taken For The Glory of God, and advancement of the Christian Faith, 
and honour of our King and Countrey, a Voyage to plant the first Colony in the Northerne 
parts of VIRGINIA, doe by these presents solemnly & mutually in the presence of God, 
and one of another, covenant, and combine our selves together into a Civill Body Politike, 
for our better ordering and preservation, and furtherance of the ends aforesaid; and by 
vertue hereof to enact, constitute and frame such just and equall Lawes, Ordinances, 
acts, constitutions, offices from time to time, as shall be thought most meet and convenient 
for the generall good of the Colony: unto which we promise all due submission and obedi¬ 
ence. 

In witnesse whereof we have here-under subscribed our names, Cape Cod, 11 of Novem¬ 
ber in the yeare of the raiyne of our soveraigne Lord King lames, of England, France and 
Ireland 18, and of Scotland 54. Anno Domini 1620. 


Mr. John Carver 

No. in family 

8 

John Turner 

No. in family 

3 

Mr. William Bradford 

2 

Francis Eaton 

3- 

Mr. Edward Winslow 

5 

James Chilton 

3 

Mr. William Brewster 

6 

John Crackston 

2 

Mr. Isaac Allerton 

6 

John Billington 

4 

Capt. Miles Standish 

2 

Moses Fletcher 

1 

John Alden 

1 

John Goodman 

1 

Mr. Samuel Fuller 

2 

Degory Priest 

1 ' 

Mr. Christopher Martin 

4 

Thomas Williams 

1 

Mr. William Mullins 

5 

Gilbert Winslow 


Mr. William White 

5 

Edmund Margeson 

1 

Mr. Richard Warren 

1 

Peter Brown 

1 

John Howland 


Richard Britterige 

1 

Mr. Stephen Hopkins 

8 

George Soule 


Edward Tilley 

4 

Richard Clarke 

1 

John Tilley 

3 

Richard Gardiner 

1 

• Francis Cook 

2 

John Allerton 

1 

Thomas Rogers 

2 

Thomas English 

1 

Thomas Tinker 

3 

Edward Doten 


John Ridgdale 

2 

Edward Leister 


Edward Fuller 

3 




More than one-half of the company perished during the succeeding months 
and descent has been proven from but the following, namely: 


1. John Alden 1 2 3 

2. Isaac Allerton 4 

3. John Billington 4 

4. William Bradford 

5. William Brewster 4 


6. Peter Brown 

7. James Chilton 4 5 

8. Francis Cooke 4 

9. Edward Doty 

10. Francis Eaton 4 


3 Includes descent from William and Alice Mullins and their daughter Priscilla. 

4 Includes descent from wife or child on the “Mayflower.” 

6 Includes descent from John Tilley and his daughter Elizabeth. 





THE MAYFLOWER PILGRIMS 


331 


11. Edward Fuller 4 


18. Henry Samson 

19. George Soule 


12. Dr. Samuel Fuller 

13. Stephen Hopkins 4 


20. Myles Standish 

21. Richard Warren 

22. William White 4 

23. Edward Winslow 


14. John Howland 5 

15. Richard More 

16. Degory Priest 

17. Thomas Rogers 4 


In contravention of the statement sometimes made, that the Mayflower 
Pilgrims were of the poorer and illiterate types of the English of that period, 
and of low social condition, the following are known to have been entitled to 
coat armor: 


John Alden 

Isaac Allerton, John Allerton 
William Bradford 
William Brewster 
Francis Cooke 
Edward Doty 
Stephen Hopkins 
John Howland 
Myles Standish 
Richard Warren 

Edward Winslow, Gilbert Winslow 


Ruskin says: “There is no wealth but life,” and Tille, Forel and Saleeby 
tell us that “a nation is composed not of property nor of provinces, but of 


men. 


In the interest of applied genealogy in relation to eugenics, a brief survey 
of the records of descendants of the Pilgrims shows that their names appear 
in places of highest honor and distinction in the history of America. Al¬ 
though not at all an exhaustive study of the subject, the following will 
indicate the value of the Mayflower spirit: 


STATESMEN 


Lineage 


John Adams, 

President United States 
Member of Congress 
Minister to England 


son of John Adams and Susan Boylston 
Joseph Adams and Hannah Bass 
John Bass and Ruth Alden 
John Alden 6 and Priscilla Mullins 8 
William Mullins 6 and Alice? 


John Quincy Adams 
President United States 
Member of Congress 


son of John Adams and Abigail Smith 
John Adams and Susan Boylston 
(See above) 


6 Passenger in the Mayflower. 




332 


SARAH LOUISE KIMBALL 


Minister to The Hague 
Minister to England 
Minister to Portugal 
Minister to Prussia 
Minister to Russia 


Charles Francis Adams, 
Member of Congress 
Minister to England 


son of John Quincy Adams and Louisa C. Johnson 
(.See above ) 


William Bradford 6 
Governor of Plymouth Colony 

John Carver 6 

Governor of Plymouth Colony 


Henry Dearborn, 
Secretary of War 
Minister to Portugal 


James Abram Garfield, 
President United States 


Ulysses Simpson Grant, 
President United States 


Frederick Dent Grant, 
Minister to Austria 

Milton S. Latham, 
Governor of California 
Member of Congress 


son of Simon Dearborn and Sarah Marston 
John Dearborn and Abigail Batchelder 
Nath’l. Batchelder and Deborah Smith 
John Smith, Jr. and Deborah Howland 
John Howland 6 and Elizabeth Tilley 
John Tilley 6 

son of Abram Garfield and Eliza Ballou 

James Ballou and Mehitabel Ingalls 
Henry Ingalls and Sibyl Carpenter 
Jotham Carpenter and Mehitabel Thompson 
Jotham Carpenter and Desire Martin 
John Martin and Mercy Billington 
Francis Billington and Christine Penn 
John Billington 6 and Ellen 6 

son of Jesse Root Grant and Harriet Simpson 
Noah Grant and Anna Buell 
Noah Grant and Susanna Delano 
Jonathan Delano and Amy Hatch 
Jonathan Delano and Mercy Warren 
Nathaniel Warren and Sarah Walker 
Richard Warren 6 

son of Ulysses Simpson Grant and Julia Dent 
(See above ) 

son of Bela Latham and Juliana H. S. Sterrett 
(1) Arthur Latham and Mary Port 

Nehemiah Latham and Lucy Harris {See) 
Arthur Latham and Alice Allen 
Chilton Latham and Susanna Kingman 


THE MAYFLOWER PILGRIMS 


333 


Robert Latham and Susanna Winslow 
John Winslow and Mary Chilton 6 
James Chilton 6 and Mary 6 

(2) Nehemiah Latham and Lucy Harris 
Arthur Harris and Mehitabel Rickard 
Isaac Harris and Jane Cooke (See) 

Isaac Harris and Mercy Latham 
Robert Latham and Susanna Winslow 

(See above) 

(3) Isaac Harris and Jane Cooke 

Caleb Cooke and Jane. 

Jacob Cooke and Damaris Hopkins 
Francis Cooke 6 

(4) Jacob Cooke and Damaris Hopkins 
Stephen Hopkins 6 and Elizabeth 6 

(5) Arthur Harris and Mehitabel Rickard 
Samuel Rickard and Rebecca Snow 
William Snow and Rebecca Browne 
Peter Browne 6 

Levi Parsons Morton, son of Daniel O. Morton and. 

Vice President United States Levi Morton and. 

Governor of New York Ebenezer Morton and. 

Minister to France Ebenezer Morton and. 

John Morton and Mary Ring 
Andrew Ring and Deborah Hopkins 
Stephen Hopkins 6 and Elizabeth 8 

Richmond Pearson son of Richmond M. Pearson and. 

Envoy Extraordinary and .Pearson and Elizabeth Mumford 

Minister Plenipotentiary at Robinson Mumford and. 

Teheran, Persia .Mumford and Sarah Christophers 

Richard Christophers and. 

Richard Christophers and Lucretia Bradley 
Peter Bradley and Elizabeth Brewster 
Jonathan Brewster and Lucretia Oldham 
William Brewster 6 and Mary 6 

son of Harrison Root and Caroline Holland 
Park Holland and Esther Bridgman 
Luther Holland and Betsey Spooner 
Philip Spooner and Elizabeth Winslow 
Kenelm Winslow and Elizabeth Clapp 
Samuel Winslow and Mercy King 
Kenelm Winslow and Mercy Worden 
Kenelm Winslow and Eleanor (Newton) Adams 
(brother of Edward Winslow 6 ) 


Elihu Root, 
Secretary of State 












334 


SARAH LOUISE KIMBALL 


Alphonso Taft, 
Attorney-General 
Secretary of War 
Minister to Russia 
Minister to Austria-Hungary 


son of Peter Rawson Taft and Sylvia Howard (See) 
(1) Aaron Taft and Rhoda Rawson 
Abner Rawson and Mary Allen 
Edmund Rawson and Elizabeth Howard 
John Howard and Sarah Mitchell 
Experience Mitchell and Jane Cooke 
Francis Cooke 6 

(2) Peter Rawson Taft and Sylvia Howard 

Levi Howard and. 

Benjamin Howard and. 

Benjamin Howard and. 

John Howard and Sarah Mitchell 
(See above) 


William Howard Taft, son of Alphonso Taft and Louisa M. Torrey 

Solicitor-General Peter Rawson Taft and Sylvia Howard 

Secretary of War (See above) 

Governor of the Philippines 
President United States 
Chief Justice Supreme Court 


Zachary Taylor, son of Richard Taylor and. 

President United States Zachary Taylor and Elizabeth Lee 

Hancock Lee, Sr. and Sarah Elizabeth Allerton 

Isaac Allerton and. 

Isaac Allerton 6 and Fear Brewster 
William Brewster 6 and Mary 6 

son of Israel Washburn and Martha Benjamin 

Israel Washburn and. 

Israel Washburn and. 

Israel Washburn and. 

Samuel Washburn and Elizabeth Mitchell 
Experience Mitchell and Jane Cooke 
Francis Cooke? 

Charles Ames Washburn, son of Israel Washburn and Martha Benjamin 
Minister to Paraguay (See above) 

Elihu Benjamin Washburn, son of Israel Washburn and Martha Benjamin 
Member of Congress (See above) 

Secretary of State 
Minister to France 


Cadwallader Colden Washburn, 
Governor of Wisconsin 
Member of Congress 


Emory Washburn, a descendant of Francis Cooke? 
Governor of Massachusetts 

Daniel Webster, 

Secretary of State 


son of Ebenezer Webster and Abigail Eastman 

Ebenezer Webster and Susanna Batchelder 










THE MAYFLOWER PILGRIMS 


335 


Benjamin Batchelder and Susanna Page 
Nathaniel Batchelder and Deborah Smith 
John Smith, Jr., and Deborah Howland 
John Howland 6 and Elizabeth Tilley 
John Tilley* 


Edward Winslow 6 
Governor of Plymouth Colony 

Josiah Winslow 6 
Governor of Plymouth Colony 

of Charles J. Wood and Caroline E. Hager 
Leonard Wood and Malvina Fitzalan Reed 
Noah Reed and Susannah White 
Micah Reed and Deborah Tomson 
Daniel Reed and Ruth White 
William Reed and Alice Nash 
William Reed and Esther Thomson 
John Thomson and Mary Cooke 
Francis Cooke* 

William Brewster* 

Stephen Hopkins 6 
William White* 

CHIEF JUSTICES 

Sir John Singleton Copley, Jr., 

Baron Lyndhurst, son of John Singleton Copley and Susanna Farnum 

Lord High Chancellor of Clarke 

England Richard Clarke and Elizabeth Winslow 

Edward Winslow and Eliz. Pemberton 
a descendant of 

John Winslow and Mary Chilton 6 
James Chilton 6 and Mary* 

son of Frederick Augustus Fuller and Catherine Martin 
Cony 

Henry Weld Fuller and Esther Gould 
Caleb Fuller and Hannah Weld 
Young Fuller and Jerusha Beebe 
Matthew Fuller and Patience Young 
Samuel Fuller, Jr., and Anna Fuller 
Samuel Fuller and Jane Lathrop 
Edward Fuller 6 and Anne* 

William Howard Taft (see Statesmen) 

U. S. Supreme Court 


Melville Weston Fuller, 
U. S. Supreme Court 


Leonard Wood, son 

Governor of Cuba 
Governor Philippines 


Also a descendant of 


336 


SARAH LOUISE KIMBALL 


SIGNERS OF THE DECLARATION OF INDEPENDENCE 

Lineage 

son of Thomas Clark and. 

Thomas Clark and. 

Richard Clark and. 

Richard Clark and. 

Richard Clarke? 

Robert Treat Paine, son of Rev. Thomas Paine and. 

Member of Continental Con- James Paine and. 

gress Thomas Paine and Mary Snow 

Nicholas Snow and Constance Hopkins? 
Stephen Hopkins and (1) ...... 

LITERATURE 

Charles Edward Banks, M.D., son of 
Banks Family of Maine (1) Edward Prince Banks and Ellen Soule 

Martha’s Vineyard, Etc. Charles Soule and Phebe Bartol (See) 

Moses Soule and. 

Moses Soule and. 

Barnabas Soule and Jane Bradbury 
Moses Soule and Mercy Southworth 
Edward Southworth and Mary Pabodie 
William Pabodie and Elizabeth Alden 
John Alden 6 and Priscilla Mullins? 
William Mullins 6 and Alice? 

(2) Charles Soule and Phebe Bartol 
George Bartol and Jane Soule 
Barnabas Soule and Jane Bradbury 
(See above) 

George Ernest Bowman, a descendant of William Brewster* 

The Mayflower Descendant 

William Bradford 6 
The Log of the Mayflower 

William Brewster, 8 
The Mayflower Compact 

son of Peter Bryant and Sarah Snell 

Philip Bryant and Silence Howard 
Abiel Howard and Silence Washburn 

Nehemiah Washburn and. 

John Washburn and Elizabeth Mitchell 
Experience Mitchell and Jane Cooke 
Francis Cooke? 


William Cullen Bryant, 
Poet 


Abraham Clarke, 
Member of Congress 












THE MAYFLOWER PILGRIMS 


337 


Winston Churchill, 
Richard Carvel 
The Crisis 
The Crossing, etc. 


Fanny Jane Crosby, 
Hymn writer 


William Howard Doane, 
Hymns and hymnals 

Ralph Waldo Emerson, 
Essayist, poet 


Josiah Gilbert Holland, 
Springfield Republican 
Scribner’s, Etc. 


son of 

(1) Edward Spalding Churchill and Emma Bell 

Blaine 

Edwin Churchill and Mary Phipps Carter 
James C. Churchill and Eliza W. Osborne 
Thomas Churchill and Alice Creighton 
Thomas Churchill and Mary Stuart Ewes 
Barnaba s Churchill and Lydia Harlow 
William Harlow and Lydia Cushman (See) 
William Harlow and Rebecca Bartlett 
Robert Bartlett and Mary Warren 
Richard Warren 6 

(2) William Harlow and Lydia Cushman 
Thomas Cushman and Mary Allerton 6 
Isaac Allerton 6 and Mary Norris 6 

daughter of John Crosby and Mercy Crosby, dau. of 
Sylvanus Crosby and Eunice Paddock 
Isaac Crosby and Mercy Foster 
Eleazer Crosby and Patience Freeman 
John Freeman and Mercy Prence 
Thomas Prence and Patience Brewster 
William Brewster 6 and Mary 6 

a descendant of William Brewster 6 


son of William Emerson and Ruth Haskins 
John Haskins and Hannah Upham 
Pliineas Upham and Hannah Waite 
Joseph Waite and Lydia Sargent 
John Sargent and Lydia Chipman 
John Chipman and Hope Howland 
John Howland 6 and Elizabeth Tilley 
John Tilley 6 

son of Harrison Holland and Anna Gilbert 
Luther Holland and Betsey Spooner 
Philip Spooner and Elizabeth Winslow 
Kenelm Winslow and Elizabeth Clapp 
Samuel Winslow and Mercy King ' 

Kenelm Winslow and Mercy Worden 
Kenelm Winslow and Eleanor (Newton) Adams 
(brother of Edward Winslow 6 ) 


Henry Wadsworth Longfellow, son of 

Poet (1) Stephen Longfellow and Zilpah Wadsworth 

Peleg Wadsworth and Elizabeth Bartlett (See) 


338 


SARAH LOUISE KIMBALL 


Percy MacKaye, 
Playwright 

Frank Munsey, 
Author, publisher 

John Howard Paine, 
Home, Sweet Home 


Benjamin Franklin Soule, 
Annals of San Francisco 


Mrs. Mercy Otis Warren, 
Historian 


Peleg Wadsworth and Susannah Samson 
John Samson (See) and Priscilla Bartlett 
Benjamin Bartlett and Ruth Pabodie (See) 
Benjamin Bartlett and Sarah Brewster 
Love Brewster 6 and Sarah Collier 
William Brewster 6 and Mary 6 

(2) Peleg Wadsworth and Elizabeth Bartlett 
Nathaniel Bartlett and Zenobe Wadsworth 

Ebenezer Bartlett and. 

Ebenezer Bartlett and. 

Benjamin Bartlett and Sarah Brewster 

(See above) 

(3) John Samson and Priscilla Bartlett 

Stephen Samson and. 

Henry Samson 6 

(4) Benjamin Bartlett and Ruth Pabodie 
William Pabodie and Elizabeth Alden 
John Alden 6 and Priscilla Mullins? 

William Mullins 6 and Alice 6 

(5) Benjamin Bartlett and Sarah Brewster 
Robert Bartlett and Mary Warren 
Richard Warren 6 

a descendant of William Bradford? 


a descendant of Stephen Hopkins 8 


son of William Paine and. 

William Paine and. 

John Paine and. 

Thomas Paine and Mary Snow 
Nicholas Snow and Constance Hopkins * 
Stephen Hopkins and (1). 

son of Charles Soule and Anne Kendall 
Moses Soule and Nancy Hewes 
Barnabas Soule and Jane Bradbury 
Moses Soule and Mercy Southworth (See) 
John Soule and Rebecca Simmons 
George Soule? 

(2) Moses Soule and Mercy Southworth 
Edward Southworth and Mary Pabodie 
William Pabodie and Elizabeth Alden 
John Alden 6 and Priscilla Mullins? 
William Mullins 6 and Alice? 

daughter of James Otis and Mary Allyn 
Joseph Allyn and Mary Doty 









THE MAYFLOWER PILGRIMS 


339 


Edward Doty and Sarah Faunce 
Edward Doty 6 and Faith Clarke 

Noah Webster, son of Noah Webster and Mercy Steele 

Lexicographer Eliphalet Steele and Catharine Marshfield 

Samuel Steele and Mercy Bradford 

William Bradford and. 

William Bradford and Alice Richards 
William Bradford 6 and Alice Carpenter 

From the foregoing records it will be plainly seen that the ideals of the 
Mayflower Pilgrims—freedom of the soul, freedom of the mind, the germ of 
our free institutions—have become a part of our national life. 

Daniel Webster, the gifted statesman and orator, a descendant of the 
Pilgrims, in an address on the first settlement of New England causes 
the Pilgrims when they landed upon Plymouth Rock to use the following 
language: 

If God prosper us, we shall here begin a work which shall last for ages; we shall plant 
here a new society, in the principles of the fullest liberty and the purest religion; we shall 
subdue this wilderness which is before us; we shall fill this region of the great continent, 
which stretches almost from pole to pole, with civilization and Christianity; the temples 
of the true God shall rise where now ascends the smoke of idolatrous sacrifice; fields and 
gardens, the flowers of summer and the waving and golden harvest of autumn, shall 
spread over a thousand valleys, never yet, since the creation, reclaimed to the use of civi¬ 
lized man. We shall whiten this coast with the canvas of a prosperous commerce; we shall 
stud the long and winding shore with a hundred cities. That which we sow in weakness 
shall be raised in strength. From our sincere, but houseless worship, there shall spring 
splendid temples to God’s goodness; from the simplicity of our social union, there shall 
arise wise and politic institutions of government, full of the liberty which we ourselves 
bring and breathe; from our zeal for learning, institutions shall spring which shall scatter 
the light of knowledge throughout the land, and, in time, paying back where they have 
borrowed, shall contribute their part to the great aggregate of human knowledge; and our 
descendants, through all generations, shall look back to this spot and to this hour, with 
unabated affection and regard. 

From Governor Bradford’s “Log of the Mayflower” down to The 
Mayflower Descendant,” by George Ernest Bowman, the records of the 
Pilgrims have been carefully preserved, covering a period of three hundred 
years. 

Many other distinguished names might be added to the foregoing list of 
eminent Americans tracing their ancestry to the Mayflower, but those men¬ 
tioned are sufficient to show that the spirit of freedom brought to our shores 
in 1620 by the founders of Plymouth Colony has been a determining factor 
in our national development, leading all endeavors along a high, spiritual 
path and bringing to our beloved country its present full measure of liberty, 
wealth and happiness. 



THE LEARNED BLACKSMITH—AN ARISTOGENIC TYPE 


HOWARD J. BANKER 

Eugenics Record Office, Cold Spring Harbor, New York 

Elihu Burritt was long known as the “Learned Blacksmith.” He ac¬ 
quired a knowledge of over thirty languages, some say fifty (5), while 
laboring daily at the forge. With slight educational advantages, his par¬ 
ents poor and the family believed to be obscure and of mediocre ability, 
many have regarded him as an example of what may be accomplished by 
the most unfavored if one but tries and perseveres. Others consider him 
an inexplicable, spontaneous genius, or human sport. Such instances are 
often cited as evidence of the impotence of heredity. 

The biographers of Burritt, who draw nearly all their material from his 
own writings, give little information concerning his family. Like many 
another genius Burritt saw his attainments as only the result of close ap¬ 
plication and determined perseverance which he believed possible to any one 
of ordinary intelligence and reasonably good health. His first public essay 
was to prove that there was no native genius. 

Without taking space to array the evidences from his life and writings 
we may briefly enumerate his most striking characteristics. He was a man 
of remarkable energy, active and restless, both physically and mentally. 
He was intensely hyperkinetic. As a boy he read all the historical and bio¬ 
graphical books in the village library of a few hundred volumes (3, 5). He 
worked at the forge twelve and fourteen hours a day and found time for 
study (3). He carried a Greek grammar in the crown of his hat that he 
might con the verbs while heating his iron (1). He left his forge in order to 
study and taught for a time. The change affected his health and he became 
a commercial traveller. He then started a grocery, failed, and walked from 
New Britain, Conn., to Boston intending to go to Europe in pursuit of his 
language studies, missed the boat and went to Worcester, returned to the 
forge and studied at the library of the Antiquarian Society, attracted at¬ 
tention by his linguistic powers, became known as the “Learned Black¬ 
smith,” began lecturing and edited a paper, advocated anti-slavery and 
temperance but chiefly international peace, travelled in Europe for three 
years, then over the United States and back to Europe promoting the peace 

340 


AN ARISTO GENIC TYPE 


341 


cause, returned to his home in New Britain at the age of fifty “without 
resources” and operated a small stony farm. He organized an Agricultural 
Club and during the Civil War returned to England and travelled on foot 
from London to John O’Groat’s studying agriculture, took other long foot 
tours (1), and so I might continue to show the restless energy of the man, 
the wanderlust and the changing activities. Altogether he published some 
thirty books on a great variety of topics besides a great many contributions 
to journals (2). 

It can not be said that Elihu Burrett was a profound, and it is doubtful 
if he was an original, thinker. He was by no means a Leibnitz. He had a 
lively intellect and great facility of expression. While laboring at the forge 
his mind demanded active exercise but there is no evidence that it ever 
took an inventive turn. He wrote books of religious meditation (1) but 
nothing to satisfy the philosophical questionings of the human soul. He 
wrote grammars for beginners in Sanskrit, Hindustanee, Persian and Turk¬ 
ish (5) but his vast fund of languages resulted in no great comparative 
philology. Great and wonderful as the man was he appears to have had 
definite limitations and his abilities appear to have been simple in character 
but extraordinary in extent. 

He unquestionably had a remarkable verbal or symbolic memory. When 
he first began work at the forge before he engaged in the study of language, 
to relieve the mental restlessness, he undertook successfully the purely men¬ 
tal calculation of such problems as: “How many yards of cloth, three feet 
in width, cut into strips an inch wide and allowing half an inch at each end 
for the lap, would it require to reach from the center of the sun to the cen¬ 
ter of the earth, and how much would it all cost at a shilling a yard?” (1) 
He could recite whole cantos of Scott and Moore (4). His memoiy doubt¬ 
less did him great service in his linguistic studies. 

He was benevolent and an idealist. The most of his life was devoted to 
the cause of promoting international peace, the bulk of his writings was in 
the interest of social reform and human betterment, in his last years he 
built mission chapels and conducted Sunday Schools (2). 

Consider now the quarry whence this character was built. Elihu had 
nine sibs of whom we know almost nothing except of the eldest, Elijah, 
and very little about him. Elijah (VI. 1) became a blacksmith studying at 
the same time mathematics. His ardor in this subject induced some friends 
to assist him to enter college (3) and he was in attendance at Williams for a 
time (4). Like Elihu, who was given a similar opportunity at Harvard, he 
could not restrict himself to a college curriculum. Leaving college he went 
south, taught school and became editor of a newspaper in Milledgeville, 


HOWARD J. BANKER 



* fci B N 


a N 


Pedigree Chart 























AN ARISTOGENIC TYPE 


343 


Georgia. His attitude on the slavery question gave offence and he was 
forced to flee for his life. Returning north he established a school of some 
prominence in his native town. He was the author of a number of school 
books, his “Geography of the Heavens” passing through numerous edi¬ 
tions and being used long after his death. He eventually went to Texas as 
the head of a small colony and there died. One of his students says, “He 
was a man of great energy of character, of a commanding presence, much 
given to ostentation and display, imperious in his manner, and too fond of 
change and adventure to make the success in life which his great talents 
would otherwise have accomplished”(4). Another writer says he “was 
a man of more than ordinary talents, but somewhat erratic” (7). 

The sister, Eunice (VI. 9), after the death of her first husband, “went west 
as a teacher, under the patronage of Governor Slade,” was saved from a 
wreck on the Lakes by “her coolness and intrepidity,” and later married a 
professor in the old Chicago University (6). She may have been a person 
of intellectual activity but we know too little about her to express an opinion. 

Isaac (VI. 3) lived near his native town. A contemporary has written, 
he “makes himself useful to church and society, has been a school teacher, 
and is earnest in every good work, has a gift in prayer, exhortation, and off¬ 
hand speaking in public” (7). 

George (VI. 4) “a young man of good promise” went south and died in 
Georgia (7). 

Of the remaining sibs we know nothing but the fact of marriage and dates. 
The study of his fraternity, meager though it be, shows that Elihu was not 
alone in some of his striking characteristics. 

The father, Elihu Burritt (V. 10),says Mary Howitt, had a hard life, 
“but his troubles and difficulties never soured the milk of his human kind¬ 
ness.” His home though small and full of its own inmates sheltered “more 
benighted travellers than any other home in the town.” The halt, the 
blind, the dumb and even the friendless and idiotic pauper became uncle or 
aunt and found a place within the family circle though having no other claim 
than their misery. Many a time returning on foot from market ten miles 
away the good shoemaker would walk two or three miles out of his way to 
bring cheer to some helpless sufferer (3). The son says he was a man of 
nervous temperament, quick apprehension and vivid sympathies (3). 
Another says he was “ a man of active and speculative mind, scrupulously 
honest and moral”(6). We seem to see here the pronounced benevolence 
of the son, perhaps something of the nervous energy and perchance the 
rudiments of the foot tours of England. 


344 


HOWARD J. BANKER 


Of the father’s fraternity we have practically nothing but genealogical 
data. One sister, Naomi (V. 6), taught school before her marriage,” was 
social and intelligent, possessing a discriminating mind” (6). The father’s 
father and family (IV. 5-7 etc.) are seemingly lost in hopeless obscurity. 
We can only guess at the connection (12). The obscurity of this Burritt 
line has tinged the whole history of Elihu Burritt as it is a common fallacy 
to think of a man’s ancestry as only in the line of the name that he bears. 

His father’s mother, Eunice Wakeman (IV. 4), had no full sibs. Of her 
and her half fraternity we know nothing of importance (6, 9). Her father 
(III. 4) was a graduate of Yale (9) and had a brother (III. 2), a graduate of 
Yale and an uncle (II. 3) a graduate of Harvard (14). They do not appear 
to have been otherwise men of great prominence. The members of this 
college group are doubtless too remote to have greatly influenced Elihu 
Burritt. What they may have contributed genetically to him it is impos¬ 
sible to say but it seems reasonable to believe that they served to raise some¬ 
what the intellectual average of the Burritt line. 

The mother (V. 11) of Elihu Burritt appears to have been superior to the 
average of her social class. The son and his biographers dismiss her with 
the respectful platitude, “ She was the best friend her children had,” men¬ 
tion “ the influence of her prayers” and “ the teachings of her godly life” (3). 
One writer, however, gives an additional touch saying that “she was a devout 
Christian woman of many resources, who could turn her hand to almost any¬ 
thing” and describes the garden with its abundance of vegetables and great 
variety of flowers; the order, neatness and comfort of the home, the group 
of mulberry trees and the cultivation of the silk-worm to which she devoted 
one room of the little house (4). Another writer quaintly remarks, “She 
was equally well versed in the Bible as in the silk business” (6). 

An illuminating incident is told of the boy’s visits with his mother to the 
village library where a few books of sermons and history could be had but as 
books could be drawn only once in two months, he endeavored to persuade 
his mother to relinquish her book of sermons that he might have one more 
of history but only occasionally did he succeed (3). This is related by the 
biographer to emphasize the son’s intellectual ardor but how much does it 
also reveal the mother’s strong intellectual hunger. And it may be signifi¬ 
cant that it was not the father in whom the boy found this intellectual 
sympathy. 

The mother had no brother and but one sister who lived to maturity— 
Roxana Hinsdale (V. 14). We know little of this sister but one statement 
concerning her is enlightening. She left at her death “some interesting 
written reminiscences of her own religious experience and life” (7). That 


AN ARISTO GENIC TYPE 


345 


a New England farmer’s wife with twelve children to care for should be im¬ 
pelled to write of her spiritual experiences would indicate that she possessed 
strong mental activities. Her son wrote several extensive genealogies and a 
local history. These two sisters evidently were endowed with some intellec¬ 
tual potentialities. There were two half sisters (V. 15, 16) but nothing has 
been learned of them. 

The mother’s father, Elijah Hinsdale (IV. 12), learned the blacksmith’s 
trade but later owned a farm and engaged extensively in silkworm culture, 
receiving for many years a bounty from the state which was seeking to 
encourage this industry (6, 8). James Freeman Clark is quoted as saying 
that “ he was a remarkable man, .... a great reader, and with only 
ordinary advantages, possessed of an extraordinary fund of knowledge,” 
and adds “His wonderful love of learning and a capability of retaining it 
will undoubtedly be found to have been handed down to the Hinsdales and 
throughout the various branches of their descendants as far as it can be 
traced” (7). This prediction is in part fulfilled in Elihu Burritt. 

Of the mother’s father’s fraternity we have significant data only of Theo¬ 
dore (IV. 16). He was a graduate of Yale and ordained to the ministry. 
He was a man “of marked ability and strong will and an active man of 
affairs” (7). The town of Hinsdale, Mass., was named for him. Until his 
death at the age of eighty “he read the classics with perfect ease and could 
quote them readily” (11). His eldest daughter, Anne, “established the 
first young ladies’ school in Pittsfield, Mass.,” which she conducted for many 
years and was later a teacher in connection with her cousin, Mrs. Willard (7). 

Of the rest of this fraternity little material is at hand except genealogical 
data, but Lydia Hinsdale (IV. 19) who married Samuel Hart was the mother 
of ten children including Emma Hart Willard (V. 19) famous as an educator 
and the founder of the Emma Willard Seminary of Troy, New York, and her 
sister, Almira Lincoln-Phelps (V. 20), only a little less eminent as an educa¬ 
tor and the author of text-books highly regarded in their day (7). 

The father of this Hinsdale group was John Hinsdale (III. 10). He was 
a blacksmith and owned a farm in Berlin, Conn. He was ensign, lieutenant 
and captain in the local militia and prominent in the community. He is 
described as a “man of rare military and literary ability,” but we have little 
definite data concerning him (7). While this is suggestive it is hardly suffi¬ 
cient ground for any very positive assertions. 

In respect to Elihu Burritt’s mother’s mother Ruth Bidwell (IV. 11) 
we know little. She died at thirty-three “supposed to have been poisoned 
by a negro slave woman.” Of her family we have practically nothing but 
some scant genealogical data. Her only brother died of consumption in 
Virginia (10). 


346 


HOWARD J. BANKER 


Reviewing the family network as a whole we see in the ancestral inherit¬ 
ance of Elihu Burritt two rather distinctly differentiated types. On the 
paternal side there appears throughout a good wholesome stock of mediocre 
ability and with no great apparent potentialities. Doubtless important 
factors in the personality of the propositus are contributed from this side 
but it is difficult to identify them with certainty and to trace their distribu¬ 
tion. His absorbing benevolence and possibly something of his nervous 
energy are derived here. What else we can hardly suggest. On the mater¬ 
nal side we find a stock that is repeatedly putting forth individuals of dis¬ 
tinct intellectual activities. What combinations were essential to call these 
out we may not know but it is evident that intellectual potentialities are 
carried in the germ-plasm of the maternal network. In Elihu Burritt this 
intellectual capacity found its expression through a combination of factors 
that gave rise to the unique accomplishments of the “Learned Blacksmith.” 
He was not the spontaneous product of a barren soil, the extraordinary 
fruition of a sterile heredity; on the contrary he was in large degree if not 
wholly the outgrowth of interlacing hereditary streams rich in the essential 
elements that constituted his personality. 

REFERENCES 

(1) Burritt, Elihu: Ten minute talks on all sorts of topics, with autobiography of 

the author. Lee and Shepard, Boston, 1874. 

(2) Northend, Charles: Elihu Burritt; a memorial volume containing a sketch of his 

life and labors with selections from his writings and lectures and extracts from 
his private journals in Europe and America. D. Appleton & Co., New 
York, [1879.1 

(3) Howitt, Mary: Memoir of Elihu Burritt. People’s Journal 2:241-246. Oct. 31, 

1846. 

(4) Lee, William H.: Reminiscences of the early life of Elihu Burritt. New York 

Genealogical and Biographical Record. 12: 101-111. July, 1881. 

(5) Camp, David N.: World’s first champion of universal peace. Recollections of 

Elihu Burritt. Connecticut Magazine 10: 599-608. 1906. 

(6) Andrews, Alfred: Memorial: Genealogy and ecclesiastical history [of New 

Britain, Conn.]: to which is added an appendix, with explanatory notes, and 
a full index. A. H. Andrews, Chicago. 1867. 

(7) Andrews, Herbert C.: Hinsdale genealogy: Descendants of Robert Hinsdale of 

Dedham, Medfield, Hadley and Deerfield with an account of the French family 
of DeHinnisdal. A. H. Andrews, Lombard, Ill., 1906. 

(8) Camp, David N.: History of New Britain, with sketches of Farmington and Berlin, 

Conn. William B. Thomson & Co., New Britain, 1889. 

(9) Wakeman, R. P.: Wakeman genealogy, 1630-1899. Being a history of the descend¬ 

ants of Samuel Wakeman of Hartford, Conn., and of John Wakeman, treasurer 
of New Haven Colony, with a few collaterals included. Journal Pub. Co., 
Meriden, Conn., 1900. 


AN ARISTOGENIC TYPE 


347 


(10) Bidwell, Edwin M.: Genealogy to the seventh generation of the Bidwell family 

in America. Joel Munsell’s Sons, Albany, N. Y., 1884. 

(11) Hawley, William A.: History of the town of Hinsdale. History of the county of 

Berkshire, Massachusetts, in two parts: the first being a general view of the 
county; the second an account of the several towns. Samuel W. Bush, Pitts¬ 
field, Mass., 1829. 

(12) Foster, Mary D.: One line of the Burritt family. West Hill Press, Ithaca, N. Y,, 

1898. 

(13) Orcutt, Samuel: History of the old town of Stratford and the city of Bridgeport, 

Conn. New Haven, 1886. 

(14) Dexter, Franklin Bowditch: Biographical sketches of the graduates of Yale 

College with annals of the college history. New Haven, 1912. 


BREVE RESENA GENEALOGICA DE LA FAMILIA IZQUIERDO 
J. JOAQUIN IZQUIERDO 

Despues de largas y pacientes investigaciones he logrado llegar a reunir 
una serie de datos relativos a mis antecesores, que me han servido para 
formar el arbol genealogico de mi familia, que va adjunto, asi como otras 
muchas noticias que espero reunir en un proximo libro. Los felices resul- 
tados de mi investigacion son enteramente nuevos, pues, con exception de 
lo relativo a las tres ultimas generaciones, los demas datos que presento 
eran totalmente desconocidos para las personas de mi familia, especialmente 
los que se refieren a las generaciones espanolas. El unico recuerdo, vago y 
borroso, conservado por algunas de las personas de mayor edad que aun 
viven, era el de que alguno de los antecesores, oyendo que en cierta ocasion 
sus hijos hacian tal o cual comentario sobre ciertos papeles nobiliarios de 
familia, que el guardaba, tras de reprenderlos asperamente por poner tanto 
interes en cosas tan mundanas, habia entregado los papeles al fuego. Ver- 
dadero o falso este hecho, lo cierto es que mi familia no poseia ninguno de 
los datos que hoy ofrezco. Si a pesar de mi espiritu liberal y propenso a 
mirar hacia el porvenir mas bien que a vivir queriendo alentar tan solo por 
el impulso del pasado, hago esta investigacion y pongo en ella cierto carino, 
es porque a mas del interes historico que creo encontrarle, pienso que bien 
puede aplicarse a su origen lo que Cesar Cantu dijo de la nobleza portuguesa, 
derivada en gran parte, como en el caso de que nos vamos a ocupar, de los 
que habian combatido en la famosa batalla de Ourique: “que no se apoyo 
en la conquista ni en el feudalismo, sino en las cualidades personales, en el 
valor, en la leal tad, en la religion.” 1 Por eso fue que, cuando depues, 
libre ya Portugal de toda domination extrangera en virtud de la decision 
del Papa Alejandro III que concedia a Alfonso Enriquez el titulo de Rey, 
que le disputaba el Rey de Castilla, y todas las tierras que pudiera quitar a 
los Moros, al reunirse las cortes de Lamego, en las que estaban representados 
el alto clero, la nobleza y los diputados de las diez y seis ciudades principals 
del reino, despues de sancionar la election que habia hecho el ejercito de 
Alonso Entiquez, por su Rey, y de coronarlo, al proceder a formar sus leyes 
y contar entre ellas las relativas al modo de componer la nobleza, las apro- 

1 C6sar Cantfi. Historia Universal. Cap. XIX, pigs. 155y 157 del tomo XXII de 
la edicidn espanola de Gass6 Hermanos, Barcelona. 

348 


GENEALOGICA DE LA FAMILIA IZQUIERDO 


349 



■Jfo, 






& J-J 


(GwTMttO 

^ i* 

_f?taiXia 'J jjtttftafi 

t X*4?* 


X>t. «$fr 


^Ru*tn!>rJ HNi.u it la \5kjt^rax»r*^u}tia< 41 ?t 3tU^i(*'.»*> 

•; it' a ^rfO- j Xttfam* 2G.\i'f. 

•*< hv-tajii utcrttf#^%k ^piaU^U ^ 
c«U ^*rntJ« J&MiCitu. 

,'ifi &uii>4.Vr $t L 31i*xt«aua Jf *t»\ 


$*$ V 'fficffixx. 


Arbor Genealogico 





350 


J. JOAQUIN IZQUIERDO 


baron “porque les parecieron buenas yjustas, doble condicion sobradamente 
olvidada en tiempos mas cultos y refinados” (Cantu). 

ORIGEN 

En los tiempos en que Alfonso Enriquez, hi jo de Enrique de Borgona y 
de Teresa, hija del Rey Alfonso I de Portugal a quien aquel, en union de 
otros caballeros franceses habia ido a socorrer, dominaba con el tftulo de 
Duque el pais que se extiende entre el Mino, el Duero y Tras-Os-Montes, y 
que de Porto-Cale, antigua capital de los gallegos, fue llamado Portugal, 
refiere Fray Jaime Bleda 2 en su “Coronica de los Moros de Espana,” que 
“ en el ano de mil y ciento treynta y nueve, el Moro Ismael mato al Rey de 
Badajoz, y se alfo con aquel Reyno, y juntando vn buen exercito con ayuda 
de otros caudillos Moros, entro en Portugal: mas el Duque Don Alonso 
Enriquez le salio al encuentro, y en el campo de Ourique, cerca de Castro 
Verde, le dio batalla y le vencio: y el Moro boluio desbaratado a Badajoz.” 
Y anade “ que esto cuentan con esta brevedad las historias arabescas, pero 
que las portuguesas dicen que el Duque salio de la ciudad de Coimbre, y 
atravesando el rio Tajo, comengo a hazer guerra a los Moros, y Ismael y 
otros quatro caudillos Moros, salieron a la defensa de sus tierras contra el 
Duque. Assento el buen Principe su Real en el campo de Ourique, cerca 
de Castro Verde: donde agora llaman cabegas de los Reyes, a vista del 
exercito de los infieles y aunque los suyos quisieron escusar la batalla, 
vista la multitud de los Moros, el los animo exortandoles con muy eficaces 
razones, y con maravilloso valor y esfuerfo: conque todos seaprestaron 
para dar la batalla, con animo de morir en la defensa de la Sancta fe. Era 
este dia a veynticinco de Julio, fiesta del glorioso apostol Santiago el Mayor, 
y assi muchos se confessaron, y comulgaron, era Martes de dicho ano. 
Alii acordaron todos, de algar por Rey al Duque Don Alonso Enriquez su 
Senor por entrar en la batalla con mayor animo. Dizen las historias 
Portuguesas, que todos a bozes aclamauan: Portugal: Portugal por el Rey 
don Alfonso Enriquez y desta forma fue alfado por Rey del inclyto Reyno de 
Portugal.” Con lo cual “quedaron muy alegres los Portugueses y luego 
ordenaron los esquadrones, que en numero eran muy inferiores a los de los 
enemigos, y arremetiendo contra ellos denodadamente, y ceuandose en 
aquella barbara gente, enemiga del nombre Christiano, mataron muchis- 
simos dellos, y tomando los cinco estandartes y pendones de los Reyes 
Moros, fue glorioso vencedor el nuevo Rey don Alonso Enriquez.” 

2 Fray Jaime Bleda. Coronica || de los Moros || de Espana. Diuidida en ocho Libros 
|| Con Licencia. || En Valencia, en la Impression de Felipe Mey. || Ano 1618. 


GENEALOGICA DE LA EAMILIA IZQUIERDO 


351 


En un nobiliario de Zazo, rey de armas que colecciono mucho, existente 
en el departamento de manuscritos del Archivo Historico de Madrid, 
(tomo XVIII, pag. 838), se refiere que “Don Diego de Guipuzcoa, el ano 
de 1139; llevando en la batalla de Ourique un estandarte, peleo tan to con 
los Moros que fueron a quitarsele, en cuya defensa perdio el brazo derecho 
y peleo como pudo con el izquierdo, sin soltar el estandarte, asistiendo en 
toda la batalla hasta que se vencio por el Rey D. Alonso I de Portugal, 
por cuyo hecho le empezaron a llamar el Izquierdo, cuyo cognomen le puso 
dicho Monarca, y le hizo grandes Mercedes. Y dio heredamientos con lo 
que se retiro a su tierra que era en las Montanas de Burgos, y alii fundo 
un solar y casa en el Valle de Sedano que es el primitivo de esta familia; sus 
descendientes fundaron otra en el lugar de Alcolea situada en la Rivera del 
Rio Sinca dos leguas de Segena y cuatro de Monzon; otra en la villa de 
Carinoso sita entre Daroca y Zaragoza, otra en Extremadura, otra en Cas¬ 
tilla la Vieja, tierra de los Cameros, y otras muchas que hay en Espana. 
Los mas usan unas Armas que son de rojo; con vanda de plata en vocas de 
dragantes verdes, lampazados de oro con dos estrellas de oro, una encima 
de la vanda y otra aba jo.” 

GENERACIONES ESPANOLAS 

Vuelto ya en su solar Castellano de Burgos, el fundador de la familia se 
desvanece, manco y glorioso, en los remotos evos de su tiempo. Ninguna 
otra noticia es dable encontrar con relation a elo a sus inmediatos des¬ 
cendientes debido a que en aquellas epocas aun no existian los archivos 
parroquiales, que no empezaron a llevarse sino hasta la primera mitad del 
siglo XVI. 

Sin embargo, sabese que por los Siglos XIV 6 XV, Don Juan Izquierdo de 
Roqas, Senor de las Torres de Roqas, en el Valle de Soba,—(solo era senor de 
la Torres de Rogas y no del pueblo que, como todo el Valle de Soba, pertene- 
cia a los condestables de Castilla, Condes de Haro),—y Patrono de su Iglesia, 
era descendiente de Don Diego, segun lo atestigua la identidad de sus armas, 
solo diferentes de las de aquel, por los esmaltes de las cabezas se dragantes 
que en vez de sinoples se vuelven de oro, y de los rayos de las estrellas, que se 
precisan ser en numero de diez. 

Fue sin duda Don Juan Izquierdo de Rog:as el que escribio en derredor 
de las armas de familia, el lema heraldico que resume su historia, relatando 
en virtud de que hechos se habia hecho valer y a costa de que sacrifices 
habia logrado construir su morada: 


352 


J. JOAQUIN IZQUIERDO 


De Rodas vine rodando 

Y a Rogasllegue rodando. 
De Rodas vine rodando 

Y a Rogas Yze de nada 

Y con sangre derramada. 


Fig. 1. Armas de la familia Izquierdo 



Aunque no he podido encontrar dato alguno que permita explicar estas 
palabras, creo leer en ellas una alusion muy clara a la famosa isla de Rodas, 
de la que se apoderaron los caballeros de San Juan de Jerusalen en 1310. 
Me parece muy probable que Don Juan Izquierdo de Rojas haya estado 
entre aquellos esforzados conquistadores, y no me sorprenderia que pos- 
teriores investigaciones me llevasen a encontrarle concurriendo a la con- 
sumacion de la obra, en 1310. Y lo repito, aunque no he podido averi- 
guarlo. Es muy probable que si no en aquel gran hecho, si haya asistido 
a algun otro, pues bien claro lo dicen sus palabras: De Rodas vine rodando, 
es decir de un lugar a otro, sin fijarse en sitio determinado, hasta que llegado 
al Valle de Soba, a Roqas hace de nada y con sangre derramada, con lo cual 
quiere perpetuar en su linage los esfuerzos que hiciera en Rodas. El nom- 




























































































GENEALOGICA DE LA FAMILIA IZQUIERDO 


353 


bre mismo de Rogas parece indicarlo: No es muy elocuente la analogfa 
entre Rodas, formado del griego pobov, rosa, y Rogas, elnombre que pone a 
su casa, de igual significado castellano? 

Se ignora con quien caso Don Juan Izquierdo de Rojas, pero se sabe que 
fue padre de Don Sancho Izquierdo de Rogas, cuyas fechas de nacimiento y 
muerte no es posible conocer por la falta de archivos parroquiales. De 
Don Sancho, se sabe que caso en Villoslada de Cameros, del Reino de Cas¬ 
tilla la Vieja, en la actual provincia de Logrono con Dona Catalina Mar¬ 
tinez de Azagra y Fernandez de Velasco y que de esta union arranca la rama de 
descendientes de Don Diego de Guipuzcoa, apellidado el Izquierdo, que 
vimos mencionar a Zazo en tierras de los Cameros , de Castilla la Vieja , de 
donde derivan, segun puede verse en el arbol genealogico que he formado, 
tanto la rama de los ascendientes del autor, como la de los Lopez Izquierdo 
Montenegro, de los que proceden los actuales Lopez Montenegro, deEspana, 
y la del Baron de Maabe a que mas adelante me referire. 

En efecto, de sus hijos, Diego Izquierdo de Rogas, que caso en Villoslada 
con Francisca Sanchez Salvador , dio origen a la rama de donde mas tarde 
habfa de salir la familia mexicana (rama Izquierdo Ruiz), y por intermedio 
de su hijo Alonso, a la del Baron de Maabe; su hermano Sancho Izquierdo de 
Rogas, que caso en Villoslada con Maria Rodriguez, dio el tronco de los 
Lopez Izquierdo Montenegro. 

Tuvo por hijos, Diego Izquierdo, a Diego Izquierdo , casado con Maria 
Garcia de Olalla, y a Alonso Izquierdo, nacido el 16 de abril de 1545, que es 
el continuador de la rama del Baron de Maabe, y que hacemos figurar en el 
arbol que he formado, porque una certification de armas extendida a su 
hijo Marcos Izquierdo Montenegro, es la que nos servira para relacionar 
a estos Izquierdo con los primitivos de que ya hemos hablado. 

A partir de esta epoca ya empezamos a contar con datos mas precisos, 
porque hallandonos a principios del Siglo XVI, empiezan a llevarse los 
libros parroquiales, que en Villoslada existen a partir de los mil quinientos 
cuarenta. 

Ya hemos dicho que Don Diego Izquierdo y Sanchez Salvador, fue 
hermano de Don Alonso Izquierdo, casado con Dona Marfa de Montenegro, 
de cuyo matrimonio resulto Marcos Izquierdo Montenegro, nacido el 4 de 
mayo de 1580, primo de Diego Izquierdo Garcia de Olalla, segun lo demues- 
tra su partida de nacimiento, que dice: “a 4 de mayo de 1580, yo el lic do 
Lopez baptice a Marcos, hijo de Alonso Hizquierdo y de M ia de Montene¬ 
gro su muger, sus abuelos Di° Hizquierdo y Antonio de Montenegro, 
vecino de Vinuesa, su padrino el lic do Hizquierdo.” (folio 16, vuelta, del 
tomo V, del libro de bautizados, finados y casados de la parroquia de N. 


354 


J. JOAQUIN IZQUIERDO 


Sra. del Sagrario de la villa de Villoslada de Cameros, Obispado de Cala- 
horra y La Calzada, Provincia de Logrono, Espana). 

La descendencia de Alonso Izquierdo, hasta el Baron de Maabe, es como 
sigue: 


Don Juan Izquierdo de Rogas 


Don Sancho Izquierdo de Rofas 

X 

Catalina Martinez de Azagra, etc. 

1 

Diego Izquierdo de Rofas 

X 

1 

Francisca Sanchez Salvador 

Alonso Izquierdo 

X 

1 

1 

Maria de Montenegro 

r 

Marcos Izquierdo 

X 

1 

Maria Izquierdo de Jimenez de Enciso 

Alonso Izquierdo 

X 

1 

Margarita L6pez de Arcos Salazar 
(en Najera) 

Alonso Izquierdo 

X 

I 

Clara de Urturi y L6pez de Arcos 
Salazar 

Ignacio Izquierdo 

X 

1 

1 

Petronila de Maturana y Ruiz de 
Otazu 

r 

Narcisa Izquierdo, 

casada con Ignacio 
de Medrano, Baron 
de Maabe. 

X 

Andrea Izquierdo, 

casada con Joaquin 

L6pez Montenegro y 

Fernandez de Velasco. 


Bendiciones 1743. 


En el archivo del Senor Don Federico Velaz de Medrano y Lopez Mon¬ 
tenegro, Baron de Maabe y Caballero de Calatrava, existe el original de una 
certification de armas, que, segun copia que debo a la gentileza de dicho 
senor, dice asi: 

Yo Domingo Geronimo De Mata Criado del Rey nuestro Senor Don Phipe quarto 
deste nombre y su Rey de armas. 

Certifico y hago entera fee y credito a todos los que la presente vieren que las armas y 
blasdn del apellido y linage de Izquierdo son las que van aqui con la verdadera razdn como 
parece por los libros y copias de linages que blasonan de los solares y armas nobles de 
Espana de mi officio que en mi poder estan, de donde lo saque en la forma que sigue. 

Los de este linage de Izquierdo son naturales de las Montanas buenos hijosdalgo de don¬ 
de han salido a diferentes partes y lugares ay de ellos en Castilla la bieja y en la Rioja y 
en el Andalucia y en otras partes. Traen por armas un escudo de goles y en el una banda 
de plata con Dragantes de Oro con apartamientos de metales y en la parte de arriba 
sobre la banda un lucero de Oro de diez Rayos y en la parte de abaxo otro lucero de otros 
diez Rayos de Oro y estas son sus armas. 









GENEALOGICA DE LA EAMILIA IZQUIERDO 


355 



Fig. 2 





















356 


J. JOAQUIN IZQUIERDO 


Las Quales dichas armas y apellido estan puestas y escritas en el libro quinto de blasones 
a fojas trescientas y noventa y siete y para que de ello conste de pedimento de Marcos 
Izquierdo Montenegro Vecino de la Villa de Villoslada dl la presente firmada de mi nombre 
y sellada con el sello de mis armas que es fecha en Madrid a ocho de Mayo de mill y sets - 
cientos y veinie y ocho Anos. 

Domm Yeronimo de Mata, rubrica. 

Rey de Armas. 

[Lugar) 

{ del \ 

[sello. j 

La anterior certification hecha a un primo de Diego Izquierdo Garcia de 
Olalla, nos permite pues, comprobar que descendian de Don Diego de Gui- 
puzcoa apellidado el Izquierdo, 1°, porque las armas son iguales a las que 
describe Zazo; 2°, porque corresponden a individuos pertenecientes a la 
rama que segun el mismo Zazo fundaron en tierras de Cameros, Tambien 
demuestra la ascendencia esta Don Juan Izquierdo de Ro$as, porque en 
derredor del blason lleva escrito el lema heraldico que hace alusion al lugar 
de su senorio, las Torres de Rogas, en el Valle de Soba. 

Caso Diego Izquierdo con Elvira Ruiz , hija de Juan Ruiz, tambien 
nacida en Villoslada. Es casi seguro que este Juan Ruiz sea el mismo 
Juan Ruiz de Villoslada, que tornado de la Historia de Gaspar Garcia de 
Alarcon, cita Fernandez Duro en la Historia de la conquista de las Azores, 
como uno de los que fueron a conquistarlas en 1583, mandando tercios con 
el titulo de Maestro de Campo, que era el que les correspondia en aquella 
epoca. 

Tuvieron por hijo, entre otros, a Pedro Izquierdo, nacido en Villoslada el 
primero de agosto de 1568, y casado en la propia villa el 15 de enero de 
1602, con Maria de la Camara, tambien nacida alii, el 18 de enero de 1585. 
(De estos datos, asi como de los que siguen, relativos a mi ascendencia 
directa, tengo los certificados de las partidas de nacimiento, defuncion y 
casamiento, autorizados por el Obispado de Calahorra). 

Su hijo Francisco Izquierdo, vino al mundo en Villoslada, el 6 de abril de 
1621 y caso el 21 de mayo de 1645 con Juana de Najera, perteneciente a la 
misma familia del Venerable Sebastian de Najera, llamado de Villoslada, 
que fue Prior del Convento de San Martin de Madrid. 

En uno de sus biografos encontramos que “la nobilisima estirpe de los 
Naxeras tuvo su origen en el Rey de Navarra llamado Don Garcia de 
Naxera, hijo del Rey Don Sancho el Mayor. Caso este valeroso Principe 
con la Reina Dona Estefania y tuvieron a los dos Sanchos V y VI de este 


GENEALOGICA DE LA FAMILIA IZQUIERDO 


357 


nombre, Reyes de Navarra.” Entre otros muchos ilustres descendientes, 
cita Fray Pedro de la Asumpsion “a San Luis Rey de Francia, a Santo Do¬ 
mingo de Guzman, al Santo Rey San Fernando, a San Martin de la Huerta 
y a Don Pedro Fernandez, fundador del Orden Militar del Senor Santiago, 
amen de otros muchos, nacidos igualmente” en Villoslada de los Cameros, 
que tiene su asiento en el Reyno de Castilla la Vieja y el Ducado de Naxera, 
famosa por su antiguedad, insigne por su vecindad y trato; podria hacerse 
copioso Nobiliario de sus muchas ilustrisimas familias” (3). 

Don Pedro Izquierdo y Ndjera , nacio en Villoslada el 9 de noviembre 
de 1648 y caso con Dona Catalina Gonzalez Orduna, nacida en la misma 
villa el 30 de octubre de 1651. Es seguro que los contrayentes debieron 
tener algun parentesco, pues la partida de su bautismo dice (folio 117, 
vuelta, del tomo septimo del libro de bautizos citado) que fue bautizada 
“Catalina yja lejitima de Ju w Gonzalez de Orduna y de Juana de Najera- 
sus abuelos paternos fueron P° Gonzalez y Maria Izquierdo.” 

Pedro Gonzalez, de Villoslada, abuelo de Dona Catalina, fue Caballero 
de Santiago en 1625 y Caballerizo del Infante Cardenal; habia nacidoen 
Malaga el 7 de agosto de 1594 y era hijo del Capitan Miguel Gonzalez, de 
Villoslada, Familiar del Santo Oficio de la Inquisition y Regidor Perpetuo de 
Malaga, oriundo de Villoslada, donde estaba casado con Dona Geronima de 
Alderete. Los padres de este Capitan eran Pedro Gonzalez y Margarita 
Sanchez, de Fuencaliente. 

No he podido averiguar ni el lugar ni la fecha hel casamiento de Pedro 
Izquierdo con Catalina Gonzalez, pues ni en Villoslada ni en Pedroso ha 
podido encontrarse el acta matrimonial. De los hijos habidos en el, Don 
Juan Izquierdo Gonzdlez Orduna es el continuador de la rama directa y el 
fundador de la familia mexicana. Nacio en Villoslada el 14 de noviembre 
de 1676, segun reza su partida de nacimiento, asentada en el folio 143 
del tomo VIII del Libro de Bautizados de esa Parroquia: “En la villa de 
Villoslada, en la Parroquial della a beinti Uno de noviembre deste ano de 
mill y seisctos y setenta y seis anos, Yo Bias Lopez, presbftero, cura y 
Veneficiado de ella Bautice y Crisme a Juan, hijo lexitimo de Pedro Hiz- 
quierdo y de Cathalina Gonzalez sus padres; Sus abuelos paternos fueron 
Fran™ Hizquierdo y Juana de Najera, y maternos Juan Gonzalez y Juana 
de Najera. Fue su padrino Juan de Najera, y nazio el contenido en catorce 
del dho mes y ano dhos - Bias Lopez - ” 


358 


J. JOAQUIN IZQUIERDO 


GENERACIONES MEXICANAS 

No he logrado averiguar como y cuando llego a la Nueva Espana Don 
Juan Izquierdo Gonzalez. Hasta los 55 anos de edad, siendo Capitan, es 
cuando lo encontramos tomando posesion del cargo Rejidor Propietario de 
la Ciudad de Puebla de los Angeles. De ello da cuenta el numero 4 de la 
Gazeta de Mexico publicada por el Bachiller Arevalo Ladron de Guevara, cor- 
respondiente al mes de diciembre de 1731, entre otras noticias de Puebla: 
“Los dlas 18 y 21 tomaron possession de Regidores de esta Ciudad, el 
Capitan D. Juan Antonio Izquierdo y D. Manuel Hidalgo de Veguellina” 
(Gazetas de Mexico. Reproducidas por el Dr. N. Leon en su Bibliografia 
Mexicana del Siglo XVIII. Mexico, 1903. Pag. 293). 

En el toirio numero 42 de los libros de cabildos de la Ciudad de Puebla, 
anos de 1729-1732, a fojas 317, vuelta, esta el acta del cabildo celebrado 
el dia 18 de diciembre de 1731, convocado expresamente por Don Pedro 
de Echeverria y Orcolaga, Justicia Mayor y Teniente de Capitan General,” 
“para entrar en possession del empleo de Rexidor de esta Novilfssima 
Ciudad a Don Juan Izquierdo.” 

En dicho cabildo, Don Juan Izquierdo presen to un Real Titulo y Superior 
Despacho, expedido por Don Felipe V, en el que se refiere que Don Juan 
Izquierdo habia hecho una petition ante el Virrey Don Juan de Acuna, 
Marques de Casa Fuerte, en la siguiente forma: 

Excelentissimo Senor: D n Juan Izquierdo, vno de los diputados del Comercio de la 
Ciu d de la Puebla de los Angeles, como mejor proceda de derecho digo quen dicta Ciudad 
se hallan vacos varios oficios de Rexidores, pues aunque hay ocho existentes, son con el 

titulo de Interinarios.Esto supuesto, pongo presente a V. Ex. que para que yo pueda 

obtener el cargo de tal Rexidor, concurren en mi las cualidades y circunstancias prevenidas 
por Reales disposiciones sin que haya impedimento que pueda embaraz&rmelo, a cuia 
vista se ha de servir su Ex ft hacerme la honra de nombrarme por Rexidor del nfimero de 
dicha Ciudad en propiedad, despach&ndoseme titulo en forma, pues por lo que a mi toca 
estoy prompto a cumplir con lo que fuere de mi obligaci6n en la forma acostumbrada por 
cuia gracia ofrezco servir a su Mag d , que Dios guarde, con un mil pesos que con efecto 
enterard en la real Caja de esta Corte despach&ndoseme titulo o concedidndoseme la gracia 
sin la solemnidad de que dicho oficio saiga a pregdn, pues si llega a salir, en ese caso no ha 
de ser visto el que ha>a yo haya de exhiuir dhos un mil pesos y se ha de hauer como si no 
los hubiera ofrecido y por esto en el caso de que halla de salir a el pregdn, s61o hago pos- 
trera ael tan sdlo en seiscientos pesos remat&ndose dicho oficio por ellos con los que enton- 
ces finicamente servird a su Mag d y no con los dhos. un mil. El Motivo que tengo para la 
separacidn de estos dos casos y posturas es que no saliendo ael pregdn el oficio, a costa de 
quatro cientos pesos m£s que van, a decir de los seiscientos, logro el tiempo que se ha via 
de ocupar en las diligencias y restituirme a dicha Ciudad con la breuedad que necesito, 
haciendoseme cargo del justo Interds que a su Magestad le resulta de esto, pues en tantos 


GENEALOGICA DE LA FAMILIA IZQUIERDO 


359 


anos como hasta oi han corrido, no ha hauido persona que se aliente a hacer postura 
alguna sin embargo de estar assl ordenado y prevenido. A V. Ex a suplico assi mande, 
que es Justicia. Juro en la forma y en lo necesario, etc. Don Juan Izquierdo. 



En vista del escrito, el “ Fiscal de su Magestad” comisionado por el 
Virrey para distaminar acerca del particular, produjo su respuesta en los 
siguientes terminos: Podra V. Ex?, en virtud de sus facultades, condescen- 
der en lo que esta parte pide haciendole merced de uno de los oficios de 
Rexidor de la expresada Ciudad con la calidad de vendible y renunciable y 
despacharle el tftulo correspondiente enterando los un mil pesos y lo cor- 
respondiente ael Real derecho de media annata con la calidad de que ha de 
traer Real confirmation dentro de los cinco anos de la ley a que es obligado 
deuajo de la pena en ella impuesta declarando para quitar toda duda ael 
tiempo de la possession deuer preferir ael suplicante a todos los Rexidores 
Interinos que hubiere, en asiento, voz y voto, sobre que V. Ex a . resolvera 
lo que tuviere por mas de Justicia que pide. Mexico y Septiembre dose de 
mil setesientos y treinta y uno. Licenziado Palacios. 

En virtud de lo anterior, y de haberse enterado en las Reales Cajas los 
pagos correspondientes, sigue diciendo Don Felipe V, en su Real Titulo: 

“En cuya conformidad y atendiendo a hauerse ejecutado los enteros 
que constan de las certificaciones insertas y que en la persona de vos es 
expresado Don Juan Izquierdo concurren las partes de calidad y demas 
circunstancias necesarias para obtener el referido empleo, con acuerdo de 
dho mi Virrey he tenido a bien elegiros y nombraros como por el presente os 
elijo y nombro, por Rexidor Propietario, uno de los del numero del Cauildo 
y Ayuntamiento de la mencionada Ciudad de la Puebla de los Angeles 



360 


J. JOAQUIN IZQUIERDO 


para que como tal lo useis y exersais por todos los dias de vuestra vida en 
todos los casos y cosas anexas y concernientes a dho empleo segun y de la 
manera que lo han hecho, podido y deuido hazer los demas Rexidores que 
han sido y son del expresado Cauildo de dha Ciudad de la Puebla, gozando 
como ellos, de todas las honras, gracias, preheminencias y demas inmuni- 
dades que por Razon de al Regidor Propietario os tocan y pertenecen 
bien y cumplidamente sin que os falte cosa alguna de ello, teniendo el lugar, 
voz y voto que os corresponde, prefiriendo en este particular a los Rexidores 
que hubiere interinos en el expresado Cauildo por estar assi declarado por 
dho mi Virrey para obviar toda duda al tiempo de la possession de este 
empleo sin poner en ello embarazo ni replica para su uso y exersicio y en 
caso de que se pon ga algun impedimento, Yo desde luego os doi por ad- 
mitido y receuido para que libremente podais seruirlo, .... 
etc., etc.” 

La preferencia que establecia el Real Titulo, por cierto justamente, 
puesto que todos los demas regidores eran interinos, fue considerada por 
estos como contraria a sus intereses, lo que motivo que Don Jorge Najera 
Enciso dirijiera al Virrey, Marques de Casa Fuerte, el siguiente memorial: 
“Exsmo. Senor: Don Juan Izquierdo Rexidor Propietario del Cauildo de la 
Ciudad de la Puebla de los Angeles con la mayor veneration parezco ante 
V. Exs a . y so las protestas de Indemnidad de todos los recursos y acciones 
de su parte, Digo que en atencion a hauer servido a su Magestad con la 

cantidad de vn mill pesos, etc.y hauiendoseme librado Real 

Titulo en forma, ocurriendo a aprehender possession del referido empleo, 
se le impugna lo resuelto por dicho Real Titulo en aquel Cauildo en quanto 
a la referida preferencia a dhos Rexidores Interinarios a quien ha convocado 
fuertemente el Justicia Mayor de aquella Ciudad sin otro motivo que el de 
perjudicarme por sus fines particulares y porque assi dicho Justicia 
Mayor como los Rexidores se hallan incursos en total inouediencia de lo 
resuelto por su Magd en dho Real Titulo se ha de seruir la Justification de 
V. Ex a ., precediendo la condigna correction y pena que fuere de su superior 
agrado, mandar se lleue a puro y deuido efecto lo resuelto por dho Real 
Titulo agrauando las penas de dho Justicia Mayor y Rexidores para en el 
caso de alguna o la mas leue omision para su exito. 

Los Regidores, por su parte, presentaron otro memorial en el que ex- 
presaban “hallarse conuenidos y ajustados en que al dicho Izquierdo se le 
de el ultimo lugar despues de los que oy se hallan apossessionados deuajo de 
las protestas de quedarle al enunciado Don Juan Izquierdo sus derechos a 
saluo para usar de ellos donde y quando le conuenga.” .... El Fiscal 
informo al Virrey diciendo que “ sin embargo de que por los Instrumentos 




GENEALOGICA DE LA FAMILIA IZQUIERDO 


361 


que vnos y otros han presentado se canoniza de justa y legal la resolution 
de V. Ex a . y que su Magestad quiere que se guarde lo mandado en la ex- 
presada ley en quanto a la preferencia y que en su conformidad se deuia 
precisar a que diese possession al dho Don Juan Izquierdo con preferencia 
a todos los Interinos, imponiendoles graues penas para su cumplimiento 
por ser frivola y sin fundamento su oposicion, sin admitirles por esta causa 
renunciation y dejacion de sus oficios y que en caso de intentarla o hazerla, 
se les deua apremiar a que prosiguiesen exersitandolos; no obstante, para 
obviar enquentros, disenciones y sin perjuicio de los derechos fiscales y del 
que compete ael referido Don Juan Izquierdo, podra V. Ex a ., siendo seruido, 
condescender por ahora a dho conuenio y mandar librar el despacho cor- 
respondiente y que los referidos oficios de Rexidores Interinarios se pregonen 
para ver si ay postores y se saquen a la almoneda.” A cuyo parecer ajusto 
el Virrey su decision, con el caracter de provisional. 

Fue entonces cuando Don Pedro de Echeverrfa y Orcolaga convoco a 
Cauildo para el 18 de diciembre de 1731. Reunido que fue, los Rexidores 
emitieron sus pareceres y, “regulados los votos, de mandato de dho Senor 
Justicia Mayor, se guardo lo uotado por la mayor parte en cuya conformidad 
la dha Nouilfssima Ciudad tomo en sus manos dho Real Titulo, lo beso 
y puso sobre su cabeza y dixo que lo obedecia y obedecio con el respeto 
deuido a carta y provision de nuestro Rey y Senor natural que Dios guarde, 
y que no puede dexar de reclamar ni pasar en silencio lo siniestro del informe 
que se hizo por Don Juan Izquierdo en que se asienta auer inducido a los 
Senores Capitulares interesados en este negocio el Senor Justicia Mayor 
Presidente de este Cauildo, siendo assi que a su solicitud se deuio la mayor 
quietud que pudiera desearse para que no resultaran enconos ni malas 
voluntades en defensa del derecho y lustre de los individuos contra quienes 
resultaba la preferencia en cuya diligencia motu propio se juntaron los 
comprehendidos de cuya action resultaron los buenos oficios que experi- 
mentaron despues de hauer librado llanamente el citatorio acostumbrado 
y respecto a que assi mismo se silencio el goze de preferencia a propietario 
que ha gozado el Senor Don Nicolas de Castro sin contradiction alguna y 
no pudiendo preferir a dicho Don Juan Izquierdo a Don Miguel Ceron sin 
causarle despojo, se ocurra por parte de dhos regidores al Real y Supremo 
Consejo de las Indias a representar todo lo que ocurre, a cuyo efecto se les 
den los testimonies que pidieren y de lo que contiene el testimonio que por 
el Senor Don Manual Santerbas se ha trahfdo de lo que se practica en la 
Ciudad de Mexico entre los Regidores propietarios e Interinos sobre el 
lugar y asiento de que gozan y a dho Don Juan Izquierdo se entre en pos¬ 
session de su empleo de Rexidor a cuyo efecto, dadose Razon por Joseph de 


362 


J. JOAQUIN IZQUIERDO 


Sumaia, Portero de Cauildo de que el sussodicho esperaba en los corredores 
de Palacio, le salieron a receuir algunos de los Senores Capitulares. En 
aceptaci6n de dho empleo se le reciuio Juramento por mi el Esscrivano, que 
hizo por Dios Nuestro Senor y la Senal de la Sancta Cruz segun derecho, so 
cargo del qual prometio usar bien y fielmente de dicho oficio de Rexidor 
como deue y es obligado, etc., etc., .... a cuya obsolucion el dicho 
Juan Izquierdo dijo Si Juro y Amen. Y esto fecho, sele dio el asiento que le 
corresponde, el ultimo despues del Senor Rexidor mas moderno.” (Tornado 
del Archivo Municipal). 

En el cabildo del 2 de enero de 1732, encontramos a fojas 339 del libro 
de cauildos citado, “que la Nobilissima Ciudad dixo que nombraua y 
nombro por Obrero Mayor al referido Don Juan Izquierdo a quien encarga 
el cuidado de las canerias yelalino de las casas de sus propios y otras cossas 
necesarias, para lo qual le daua comission en bastante forma y el poder que 
de derecho se requiere y sea necesario y por el trabajo y ocupacion que en 
ello ha de tener, le asignaba y asigno cien pesos de salarios, pagados de sus 
propios como es costumbre.” 

Sin embargo, no pudo Don Juan Izquierdo llevar a cabo su obra, ni lie- 
go a saber la resolucion del Consejo de Indias, porque fallecio el dia 4 de 




GENEALOGICA DE LA FAMILIA IZQUIERDO 


363 


marzo de ese ano, segun puede verse en su partida de defuncion, que logre 
encontrar en el archivo del Sagrario de Puebla y que en facsimile va 
a continuation. 

He recorrido minuciosamente el templo de San Cristobal de Puebla, en 
donde la partida anterior declara que fue enterrado Don Juan, sin encontrar 
huella alguna de su tumba. El piso ha sido renovado, quiza no una sino 
varias veces en el espacio de tiempo transcurrido hasta la fecha, de cerca 
de doscientos anos, y es seguro que por eso la lapida tambien ha 
desaparecido. 

En la Gazeta de Mexico numero 52, correspondiente al mes de marzo de 
1732, entre las noticias de Puebla de los Angeles, sigue la de que “el dia 
4 murio el Capitan Don Juan Antonio Izquierdo, Rexidor de esta Ciudad 
y Diputado de Reales Alcabalas. (Gazeta de Mexico con licencia del 
Excmo. Senor Virrey. En Mexico, en la Imprenta Real del Superior 
Gouierno, de los Herederos de la Viuda de Miguel de Rivera Calderon, en 
el Empedradillo). 

A fojas 387 del tomo citado del archivo muncipal de Puebla, el Pro- 
curador Mayor informa de haber revisado las cuentas de Don Juan Izquierdo 
y dice: “que las tiene reconocidas con los instrumentos que se han pre- 
sentado de su comprouasion y dellos consta que hauiendo sido electo Obrero 
Mayor de esta Nouilissima Ciudad el Senor Don Juan Izquierdo su Capitular 
ya difunto, exersio dicho empleo el tiempo que uiuio desde principios de 
este ano en que fabrico y reforzo las presas del rfo que llaman de San 
Francisco, por donde passalacaneria del agua que uiene a esta dicha Ciudad 
y juntamente hizo otras obras necesarias al bien publico, en que gasto la can- 
tidad de quinientos quarenta pessos quatro reales y medio como parece de 
los reciuos que ha presentado el Cap n D n Xpoual Martinez de Castro, 
Aluasea de dho Senor difunto.” 

El Capitan y Regidor Perpetuo de la Puebla de los Angeles, Don Juan 
Izquierdo Gonzalez, en fecha que no he podido averiguar, caso con Dona 
Josef a Martinez de Castro , cuya genealogia era como sigue: 

General Gabriel Guadalajara — Marfa Ana Caballero 


Pedro Guadalajara — Maria Ana de Sagasti 


Francisco Guadalajara — Tomasa Martinez 


Gertrudis Guadalajara — Cristobal Martinez de Castro 
Josefa Martinez de Castro 





364 


J. JOAQUIN IZQUIERDO 


Ignoro sus fechas de nacimiento y de muerte y solo he podido averiguar 
que nacio en Puebla y que despues de la muerte de su esposo caso en 
segundas nupcias con el Capitan Don Nicolas Gomez, de Rucoba, tambien 
Regidor de la Ciudad de los Angeles, que fallecio el 2 de enero de 1755, 
dejandola viuda, y que esta enterrado en el templo de Santo Domingo, de 
Puebla. 

Del matrimonio de Don Juan Izquierdo y Doha Josefa, nacio Don 
Joseph Joachin Izquierdo , el 28 de diciembre de 1730, siendo bautizado el 
3 de enero del ano siguiente, en el Sagrario de la Catedral Angelopolitana, 
segun consta en su acta de bautismo, puesta a continuation (Archivo 
Parroquial del Sagrario de Puebla; libro de bautismos numero 31; anos de 
1729-1730, a fojas 245). En ella se encuentra consignado el lugar de 
origen de Don Juan Izquierdo, su padre, dato que me sirvio de punto de’ 
partida para investigar lo relativo a las generaciones espanolas. 

Don Joseph Joachin tambien fue Capitan y Rejidor Perpetuo de la 
Ciudad de Puebla. Rebuscando en el archivo municipal de la Ciudad, 
encontre que ingreso como Rejidor el 26 de junio de 1755. En el cabildo 
de ese dia se vio su Real Titulo, expedido por Fernando VI, en el cual se 
refiere como, Don Nicolas Gomez de Rucoba que segun vimos, habia 
casado en segundas nupcias con Doha Josefa Martinez de Castro, madre 
de Don Joseph Joachin, habia “renunciado el oficio de Rejidor de la No- 
bilissima Ciudad, en primer lugar en Don Joseph Joachin; en segundo, en 
Don Joseph Izquierdo, vecino y Labrador de la Provincia de Atlixco, y en 
tercero en Don Juan Enciso de Thexada.” Consta despues la siguiente 
certification: “Manuel del Castillo, Escriuano del Rey Nuestro Senor que 
Dios guarde, Publico, Propietario y uno de los del numero de esta muy noble 
y muy leal Ciudad de los Angeles, Notario Receptor de los Juzgados 
eclesiasticos deeste Obispado y del Sancto Oficio de la Inquisition de este 
Reino. Certifico y doy fee en testimonio de verdad que oy dia de la fecha, 
cosa que seran las quatro y media de la tarde, estando en las casas que f ueron 
de la morada del Cap n D n Nicolas Gomez de Rucoba Rejidor deesta Nma. 
Ciudad, veo en la Sala principal de ella el cuerpo del citado Dn. Nicolas 
amortajado con auito de nuestro Seraphico Padre Sehor San Fran co , que, 
se halla al parecer muerto y sin espiritu de vida, al qual conod, trate y 
comiinique uiuiendo y para que conste donde convenga, de pedimento de 
Dn. Joseph Joachin Izquierdo Gonzalez doy la presente en la referida ciudad 
de los Angeles a treinta y un dias del mes de Diciembre de mil setecientos 
sinquenta y quatro anos, .... hago mi signo en testimonio de verd¬ 
ad. Manuel del Castillo Escriuano Real y Publico.” 


GENEALOGICA DE LA FAMILIA IZQUIERDO 



365 


Facsimile No. 3 




366 


J. JOAQUIN IZQUIERDO 


LLenos los demas requisitos y formalidades necesarias, que aqui es im- 
posible enumerar; evaluado debidamente el oficio renunciado, por varios 
peritos; enterados en la Real Caja los pagos relativos y comprobado todo 
ello debidamente, sigue diciendo Don Fernando VI en su Real Titulo: 
“En cuya conformidad y atendiendo a que en la persona de Vos Dn Joseph 
Joachin Izquierdo Gonzalez Martinez de Castro concurren las partes de 
calidad, idoneidad, suficiencia y demas circunstancias, con acuerdo del 
repetido mi Virrey, Conde de Reuillagigedo, he tenido a bien elegiros y 
nombraros como por el presente os elijo, proueo y nombro, por Rejidor, uno 
de los del numero del Cauildo de la Ciudad de la Puebla de los Angeles para 
que como tal Rejidor propietario useis y exersais este empleo en todos los 
casos y cosas a el anexas, .... etc., etc.” 

El Capitan y Rejidor Don Joseph Enciso de Thexada, dijo: “que 
obedeciendo como obedece con el respeto y veneracion deuida al Superior 
mandato de que se ponga en posesion del Rejimiento que se hay a despachado 
Don Joachin Izquierdo, pero en quanto a su cumplimiento haze presente a 
esta Nma. Ciudad berzarse en dha posesion el interes Real a cuio aug- 
mento todos estamos obligados por estar hecha la renuncia de dho. Re¬ 
jimiento en virtud de la qual se le libraron despachos en sujeto menor de 
edad y no se le hubieran librado con noticia de serlo, pues para que fuesse 
valida necesitaua la gracia y expresa concesion de S. Magd., que Dios guarde, 
en la que a Don Nicolas Gomez de Rucoba le concediesse facultad de 
poder executar las renunciaciones del oficio de Rejidor que obtenia en sujetos 
menores de edad .... por lo que procurando el mejor y mas seguro 
acierto en el asumpto y su conciencia, es su voto se llame a este cauildo 
el abogado de esta Novma. Ciudad para que en inteligencia y con prospecto 
a los despachos presentados por el pretendiente y las leyes que tratan del 
asumpto y el interes Real, diga lo que se deua ejecutar porque las leyes lo 
tendran determinado a beneficio de su Magestad sin perjuicio de sus vasallos. 
Y en su vista dha Nouilissima Ciudad mando llamar al Licdo. Don Joachin 
Maria Infante de Zetina Abogado de la Real Audiencia de Mexico y de esta 
Nma. y auiendo uenido y entrado en la Sala Capitular se lehizo sauer el uoto 
antecedente y lo decisiuo de dho Real Titulo. Expuso su dictamen en papel 
separado cuio thenor es como sigue: “ el Asesor expuso lo siguiente, que por la 
ley Real decima del titulo veintede la Recapitulation delndias esta preuenido 
que no se hagan renunciaciones de oficios en personas que sean menores de 
edad, pena de perdimiento de los oficios y de no ser admitidas. Y que los 
exmos Senores Virreyes no puedan dispensar en tales cassos, aun a titulo de 
compassion. Pero que ael mismo tiempo se hace cargo de que la notoria 
Literatura del Senor Fiscal tuvo muy presente como siempre esta ley Real; 


GENEALOGICA DE LA FAMILIA IZQUIERDO 


367 


y que su Senoria no opuso a este defecto, o porque no le constaria ser punto 
de hecho; o porque segun se informa y se asienta en este cauildo, el prouisto 
pasa de ueinticuatro anos y seis meses, lo que junto con la hauilidad y partes 
del prouisto prestarian motiuo bastante para disimular en el corto tiempo 
que le falta para completar la edad; y que a esto se anade que en el mismo 
titulo se expresa la clausula de que no se le ponga ningun defecto; y que si 
de hecho se le opusiere, por el mismo caso, se manda apossesionar; con cuios 
fundamentos concurre tambien la exequibilidad que piden estos Reales 
Rescriptos particularmente en el caso presente; en que media la Publica 
expectacion, pues la Ciudad y Republica convidada para la possession y 
aun la pleue, a quien se ha trascendido la noticia, estan en la expectacion, 
de que resultaria desairado el Real Rescripto, los poderosos respetos que en 
el se ueneran y las circunstancias que uan prenotadas, esto mediante es de 
dictamen que sin perjuicio de lo que por la Souerama de su Exa. se deter- 
minare y de lo que por la notoria justification del Senor Fiscal se pidiere 
con noticia del impedimento por falta de edad, se le de al citado Don Joseph 
la posesion con arreglamiento al Real Titulo y que sobre el impedimento 
opuesto por defecto de la edad se consulte a la Souerama de su Exa para 
que determine como siempre lo mejor, . . . etc.” “Y por la Nov a 

Ciudad oydo el parecer antecedente dijo que conformandose como dha 
Nouilissima se conformacon el, acordaua y acordo se haga en todo como por 
dho Asesor se consul to y en su inteligencia mandaua y dha Nov a mando 
a Dn Fran co Valentin Ordonez sustituto del Portero de Cauildo pasase a 
las casas de hauitacion y morada del dho Don Joseph Joachinlzquierdoyle 
diesse razon como esta Nov a le estaua aguardando, lo que executo dho 
sustituo de Portero de Cauildo y hauiendo uuelto dio noticia a dha Nov a 
que el referido Don Joseph Joachin se hallaua en los corredores de este 
Palacio y le salieron a receuir algunos de los Senores Capitulares y hauiendo 
entrado en dha Sala Capitular por ante mi el Escriuano juro por Dios nuestro 
Senor y la Senal de la Sancta Cruz en forma de derecho, de vsar bien y 
fielmente el empleo de Rejidor, etc. etc. . . .” (Archivo Municipal, 

loc. cit.). 

El dia siete de agosto del mismo ano, presento ante el cabildo un Superior 
Despacho de Don Juan Francisco de Giiemez y Horcasitas, Conde de Re- 
uillagigedo, Gentil hombre de la Camara de Su Magestad, con entrada, 
Tenien te General de sus Reales exersitos, Virrey Gouernador y Capitan 
General de esta Nueva Espana y Presidente de la Real Audiencia de ella, 
etc., por el cual, atendiendo a que Don Jose Joaquin habia comprobado con 
su partida de bautismo, tener veinticuatro anos y cerca de siete meses, 
fundandose en que la ley 17, titulo tercero, del libro septimo de la Recapitu- 


368 


J. JOAQUIN IZQUIERDO 


lacion de Castilla solo exigia la edad de veinte anos para obtener el oficio de 
Rejidor, declaraba “por ualida la renuncia hecha por Don Nicolas Gomez 
de Rucoba en el referido Don Jose Joaquin y en su consecuencia, por sub- 
sistente la posesion que se le habia dado del oficio de tal Rejidor del numero 
de la cuidad de la Puebla por su Cauildo y Ayuntamiento a los ueintiseis de 
junio proximo pasado de este ano.” 



Facsimile No. 4 


Anticipandose a la llegada del nuevo Virrey, en el cabildo del 23 de julio 
de ese mismo ano de 1755, “la Nov a Ciudad dijo que nombraua y nombro 
a los Capitanes y Rexidores Don Juan Joachin Micieses Altamirano y 
Don Joseph Joachin Izquierdo para que fueran al Pueblo de Thepellahualco 
a Receuir al Ex m0 Senor Virrey, proximo a conducirse a esta NueuaEspana.” 
En la description del viaje de la Marquesa de las Amarillas, hecha en verso 
por Don Antonio Joaquin de Rivadeneyra Barrientos, (reimpresa con 
notas de Don Manuel Romero de Terreros y Vinent, Marques de San Fran¬ 
cisco, en los Anales del Museo Nacional, tomo V, num. 4, 1914, pag. 235), 
se refiere que el encuentro tuvo lugar el 21 de octubre, segun rezan, los 
siguientes versos, ramplones y de mal gusto como todos los de la obra: 

A Perote venimos, 

distante quatro leguas, donde hicimos 
mansi6n la noche, siendo la jornada 
a Poblaci6n nombrada 
Thepeyahualco la del dia veinte y uno. 

Alii nos alcanzo uno 

de los dos Secretaries de Gouierno, 


GENEALOGICA DE LA PAMILIA IZQUIERDO 


369 


y con las muestras de un amor paterno, 

Augustin recibio los Diputados, 
que llegaron embiados, 
de la Villa y la Puebla. 

La ciudad se aprestaba a recibir a sus huespedes y los comisionados 
para su recibimiento, con el fin de disponer los alojamientos, presentaron 
la siguiente curiosa relacion de la f am ilia que traia el Maques de la 
Amarillas, que reproduzco por lo interesante y pintoresca que la encuentro: 
(fojas 149, vuelta, del tomo citado): “Razon de la familia que trae el 
Senor Virrey. = La Senora marquesa D na Luisa de Ahumada uiene en 
sinta = D n Augustin de Ahumada y Villalon su hijo de ueinte y ocho 
meses con grado de Coronel y Cap n de la Compania de Cauallos de Palacio 
= Don Manuel Baamont, etc. etc. = Damas y criados, ocho = Pajes y 
Marmitones, mas de sesenta = confesor, el padre Fray Pedro Moreno, domi- 
nico. = Trae su excelencia un ajuar muy exoruitante.” 

Las comisiones principales que desempeno Don Joseph Joachin Izquierdo, 
fueron: comisario de tierras en el ano de 1756; diputado, veedor de car- 
nicerias, patron de fiestas y procurador de pobres, en los siguientes. 

En el cabildo del 15 de junio de 1757, presen to un segundo real titulo 
de confirmation de su Regimiento, expedidopor Don Fernando VI, y despues, 
estando para llegar el Nuevo Virrey Dn. Francisco Cajigal de la Vega, fue 
nombrado para darle la bienvenida, en Tepeyahualco, pero no llego a desem- 
penar la comision por haberse excusado por motivos de enfermedad. 

En 1766, presen to a Don Joaquin de Monserrat, Marques de Cruillas, 
la “renuncia de su cargo de Rexidor, por motiuo de hallarse hauitualmente 
enfermo de hauer experimentado nosiuo el temperamento de aquella 
Ciudad. Se hauia retirado a su hazienda de labor asistiendo en ella la mayor 
parte del ano por recuperar assi su salud con el natural deseo de conservarla; 
que este impedimento fisico de asistir en dha Ciudad le era consequente- 
mente de seruir el empleo de Rexidor dexandole en el desconsuelo a que tal 
vez por su falta experimentase el Publico algun perjuicio y para euitarlo 
hauia deliberado separarse enteramente de el, y por lograrlo con utilidad 
de la Real Hazienda, en seruicio y obsequio de su Magestad, desde luego 
lo renunciaba a su fabor en mis manos para que se vendiesse a su beneficio 
con la calidad de gozar de los honores que de ninguna manera renunciaba.” 

. . . El Virrey, en su vista y teniendo en cuenta la respuesta del 

Fiscal, de fecha 12 de noviembre de ese ano, decia: “teniendo presente que 
la renuncia que a favor de su Magestad hace Don Joseph Joachin Izquierdo 

es sin duda a beneficio de su Real Auer, etc.no ai in- 

conveniente en que yo le conceda la renuncia atento el particular seruisio 


370 


J. JOAQUIN IZQUIERDO 


referido, entendiendo se de los priuilegios y excempciones que no esten 
adictos al actual exersicio y de aquellas que sin agrauio o perjuizio de tercero 
solo siruan para mantener el decoro, esplendor y estimation deuida a un 
sujeto que ha sido miembro de Aquel Ayuantamiento y se aparta de el 
por su voluntaria renuncia.” .... 

El Capitan y Rejidor Don Joseph Joachin Izquierdo, caso con Dona 
Maria Gertrudis Yanez de Vera , hija de Don Ignacio Yanez de Vera 
y de Dona Josef a Camino Frias. No he llegado a saber las fechas de naci- 
miento y defuncion de ella, ni la del matrimonio. 



Fig. 3. El Capitan y Regidor de la Puebla de los Angeles Don Joseph 
Joachin Izquierdo 

Murio Don Jose Joaquin en su hacienda de San Bias (Estado de Tlaxcala). 
el 20 de septiembre de 1779, dejando viuda a su esposa. Su cuerpo fue 
llevado al Santuario de Ocotlan, a extramuros de Tlaxcala, donde fue 
inhumado. En un libro de entierros existente en el convento de San 
Francisco de la cuidad de Tlaxcala, consta la partida siguiente: 

La familia conserva un retrato de Don Joseph Joachin, que mi distin- 
guido amigo el Marques de San Francisco describe asi: “El retrato es en 
cera, probablmente de los que ejecutaba Francisco Rodriguez a fines del 
siglo XVIII y principios del XIX. Como casi todos sus congeneres, el 
personaje esta representado de perfil y colorido. El cabello, gris, esta 
peinado a la manera de la epoca en que empezaron a desecharse las pelucas. 


GENEALOGICA DE LA FAMILIA IZQUIERDO 


371 


Viste uniforme militar azul palido con vueltas negras y ribetes rojos; la 
chupa es encarnada; y ostenta banda azul y blanca, aparentemente de 
Carlos III, pero el azul es mucho mas oscuro que el que corresponde a esta 
Orden. Como los retratos en cera no se generalizaron en Mexico sino hasta 
despues de 1779, este retrato debe haber sido hecho despues de la muerte 
del Senor Izquierdo. La caja ovalada de plata que lo contiene esta sur- 
montada por una corona del mismo metal, que evidentemente no le 
pertenece.” 




__ _ eJ- -3 <-C\ 

C««- Zy** cCt 

cXe. rrJ.LS<-l ta-j .5 nu. e. a. ->^try- 

CC l ?/ta re'. z ^ Cul ra_*. 

S *’*’ y <-0 CcxZcc. 

c. y.f'^.Oocin'n <*., /«.c/« '5tfio*Uu .1 

7^ ^ Cc / a a ‘ c. cA*. C^J<n «-Xac A/n c&e, 

,'r rCc *-6io 6*1 s Senary 

A*/>.c<V.V»o/r. 

c-^ Vc^.c 

x/x- ^ 


Facsimile No. 5 


Aunque no he podido saber que uniforme es el que lleva y tampoco han 
podido reconocerlo diferentes historiadores, amigos mfos, me parece muy 
probable que se trate del del Regimiento del Comercio de Puebla que hallo 
descrito en el Calendario Manual y guia de for aster os para el ano del Senor 
de 1787, dispuesto por Zuniga y Ontiveros. Consta allf que “ el Regimiento 
del Comercio de Puebla, esta formado de un batallon: Fue creado en 1740. 
Esta aprobado por S. M. bajo las mismas reglas y Priuilegios que el de 
Mexico. Su uniforme: casaca y calzon azul , chupa y vuelta encarnada , 
boton bianco, y los oficiales galon de plata al canto de casaca y chupa.” 

De los siete hijos de Don Joseph Joachfn y de Dona Maria Gertrudis, 
que pueden verse en el arbol, Don Jose Ignacio Mariano Izquierdo , es el 
continuador de mi rama directa. Nacio en la hacienda de San Bias el 4 de 


372 


J. JOAQUIN INQUIERDO 


julio de 1769 y caso, en fecha que ignoro, con Dona Mariana de Hogal. 
Hasta el presente no me ha sido dado averiguar la fecha y el lugar de su 
muerte, asi como los de su esposa. 

Don Joaquin Maria de JesiXs Izquierdo, su hijo, nacio en San Bias, el 16 
de mayo de 1802 y fue hermano de otros nueve que aparecen en el arbol. 
Caso en primeras nupcias con Dona Rosalia Bernal, madre de mi abuelo, 
y en segundas con Dona Cayetana Hogal, de la cual resulto una pequena 
rama que aparece en el arbol, ya extinguida. Su esposa murio el 7 de 
junio de 1844, y el, cinco anos mas tarde, el 3 de abril, jueves santo de 1849. 

De su union, nacio el senor mi abuelo, Don Manuel Izquierdo y Bernal 
en 1839, y sus hermanas Dolores (1835), Soledad (1837) y Mariana (1838), 
priora, la primera de ellas, de las monjas inesas de Puebla. 

Don Manuel Izquierdo murio en su hacienda de San Bias, el 11 de julio 
de 1889, y su esposa, Dona Trinidad del Pozo, en Puebla, el 11 de julio de 
1898. En su union, tuvieron por hijos, a Don Carlos Maria Izquierdo, 
actual propietario de la Hacienda de San Bias, la centenaria propiedad de 
la familia, nacido en 1859; a Doha Isabel, nacida en 1860; a Don Joaquin 
Pelayo, mi padre, nacido en 1865; y a Doha Maria y Don Antonio, nacidos 
en 1867 y 1873, respectivamente. 

Don Joaquin Pelayo Izquierdo , mi padre, nacio en Puebla, el 26 de ju 
nio de 1865, y caso en la misma ciudad, el 27 de abril de 1892, con Doha 
Maria Rauddn Asunsolo, nacida en Aguascalientes el 4 de diciembre de 
1866, hija de Don Ignacio Raudon, coronel que fue del Ejercito Mexicano, 
condecorado con la Cruz de la Angostura por haber combatido en esa action 
contra el invasor yankee, y caballero de la Imperial Orden de Guadalupe, 
y de Doha Emilia Asunsolo. 

Don Joaquin Izquierdo fue Rejidor de la Cuidad de Puebla, casi a los 
doscientos anos de que lo habian sido sus antecesores. Pero como los tiem- 
pos eran completamente diferentes, lo fue por election que por mucho tiempo 
sera entre nosotros la unica que verdaderamente fue democratica, el 11 de 
diciembre de 1911. Ingreso como rejidor propietario el 30 de marzo del 
ano siguiente, de 1912, ancargandose de la seccion de policia, y por varios 
dias desempeno interinemante la presidencia municipal, en octubre y no- 
viembre del propio ano. El 12 de noviembre se hizo cargo delasegunda 
seccion de hacienda, y el 31 de marzo de 1913, le era concedida una licencia 
ilimitada para separarse de la corporation municipal. Murio el 12 de 
agosto de 1913. 

Jose Joaquin Izquierdo, nacido en Puebla el 8 de mayo de 1893, hizo sus 
estudios de medico-cirujano en el Colegio del Estado de Puebla y sus ten to 
su examen profesional los dias 4, 5 y 8 de enero de 1917. A1 recibir su 


GENEALOGICA DE LA EAMILIA IZQUIERDO 


373 


titulo, vino a ser el primer profesionista habido en la familia mexicana. 
Es miembro de la Sociedad Cientifica “Antonio Alzate,” Jefe de Trabajos 
practicos de Fisiologia y Biologia en la Escuela Nacional de Medicina, 
Socio titular de la Academia Nacional de Medicina y fundador de la Sociedad 
Mexicana de Biologia, etc. 


THE ONEIDA COMMUNITY EXPERIMENT IN STIRPICULTURE 


HILDA HERRICK NOYES, M.D., and GEORGE WALLINGFORD NOYES, A.B. 

The institution afterwards known as the Oneida Community was founded 
by John Humphrey Noyes at Putney, Vermont, in 1841. In 1848 under 
pressure of persecution the Community removed to Oneida, New York, 
which at that time was a frontier settlement. In its new location the 
Community grew and prospered until January 1, 1881, when, for reasons 
which need not here be detailed, it was re-organized as a joint-stock 
company. Since that date the joint-stock successor of the Community has 
carried on its business with extraordinary success, its net assets having 
increased from about $550,000 in 1881 to nearly $7,000,000 at the present 
time. 

Noyes first published his views regarding human “ stirpiculture,” as he 
called it, in the “First Annual Report of the Oneida Community” in 
February 1849. He says: 

We are not opposed to procreation. But we are opposed to involuntary procreation. 
We are opposed to excessive and, of course, oppressive procreation, which is almost 
universal. We are opposed to random procreation, which is unavoidable in the marriage 
system. But we are in favor of intelligent, well-ordered procreation. We believe the 
time will come when scientific combination will be applied to human generation as freely 
and successfully as it is to that of other animals. 

For twenty-seven years after its foundation the Oneida Community was 
engaged in working out the immediate problems of its new order of society, 
and could not undertake an experiment in scientific propagation. But in 
1868 it found itself in a situation favorable for such an experiment in the 
following respects: 

1. A social system based on their religion, called “complex marriage,” 
which, under stringent regulations and circumscribed by the limits and 
approval of the Community, permitted the requisite flexibility in mating. 

2. Sufficient income. From 1841 until 1855 expenses exceeded income. 
After 1855 income exceeded expenses, and during the period 1868-1872 the 
net income above all expenses averaged more than $47,000 per year. 


374 


THE ONEIDA COMMUNITY EXPERIMENT 


375 


3. Sufficient number of members to admit a considerable degree of selec¬ 
tion. The membership during the eleven years of the stirpicultural experi¬ 
ment ranged from 271 to 306, with a slight excess of women over men. 

4. Capable leaders. Noyes himself was a graduate of Dartmouth 
College, had studied theology at Andover and Yale, and his management of 
the Community during twenty-seven years had fully demonstrated his 
unusual ability. He was surrounded by a cabinet of men and women 
(among them a number of college graduates), who had shown the qual¬ 
ities of leadership in positions of high responsibility. 

5. Confidence on the part of the members in their leaders and in the 
stability of the Community. The members were practically unanimous in 
believing that Noyes and his subordinate leaders were inspired and that 
the Community was the earthly representative of the Kingdom of God. 
Without this superlative confidence they would not have been willing to 
engage in such an experiment. 

6. A method of subjective, healthful, reasonably effectual birth-control 
called “male continence,” discovered by Noyes in 1844, which did not 
exclude from sexual communion those judged unfit to take part in the 
stirpicultural experiment. 

7. A high degree of passional control due to the fact that discipline and 
effort had from the beginning been constantly directed toward perfection 
of character. In the early days no little trouble had been caused by 
jealousy and exclusiveness, but by 1868 these anti-Communistic traits had 
been so far conquered that altruism deemed impossible in ordinary society 
was commonplace in the Community. 

8. Extraordinary religious devotion. This characteristic made a large 
majority of the Community members fervently desirous not alone negatively 
to avoid selfishness, but positively to engage with all their powers in any 
enterprise which they thought in the interest of the Kingdom of God. 
Such pre-eminently was the enterprise of human redemption by “combining 
regeneration with scientific generation.” 

While the Community did not pretend to unusual knowledge of stir- 
piculture, the leaders nevertheless devoted much study to the literature of 
the subject, especially the works of Charles Darwin and Francis Galton. 
Noyes embodied the results of his studies in an article entitled “Scientific 
Propagation,” which was published in The Modern Thinker , August 1870, 
and later in pamphlet form. 

Early in 1869 the young men and women of the Community made the 
following declarations: 


376 


H. H. AND G. W. NOYES 


Statement addressed to J. II. Noyes by young men 

The undersigned desire you may feel that we most heartily sympathize with your 
purposes in regard to scientific propagation, and offer ourselves to be used in forming any 
combinations that may seem to you desirable. We claim no rights. We ask no privi¬ 
leges. We desire to be servants of the truth. With a prayer that the grace of God will 
help us in this resolution, we are your true soldiers, 

(Signed by 38 young men.) 


Resolutions of young women 

1. That we do not belong to ourselves in any respect, but that we do belong first to 
God, and second to Mr. Noyes as God’s true representative. 

2. That we have no rights or personal feelings in regard to child-bearing which shall in 
the least degree oppose or embarrass him in his choice of scientific combinations. 

3. That we will put aside all envy, childishness and self-seeking, and rejoice with 
those who are chosen candidates; that we will, if necessary, become martyrs to science, 
and cheerfully resign all desire to become mothers, if for any reason Mr. Noyes deem us 
unfit material for propagation. Above all, we offer ourselves “living sacrifices” to God 
and true Communism. 

(Signed by 53 young women.) 

The experiment which followed these declarations occupied the years 
1868-1879 inclusive. About 100 men and women took part, of whom 81 
became parents. During the eleven years 58 living children were born, 
and there were 4 still-births. A precise appraisal of the factors entering 
into the experiment and of its outcome must be the work of scientific special¬ 
ists, preferably those wholly unconnected with the experiment itself. The 
present paper attempts only a brief, tentative account. 

The Oneida Community was a product of successive unconscious group 
selections. To go no further back than the English Puritans four such selec¬ 
tions can be distinguished: 

1. The New England pioneers were a selection from the English Puritans. 

2. The Revivalists were a selection from the New England pioneers. 

3. The Perfectionists, comprising those who believed in the possibility 
and obligation of freedom from sin in this world, were a selection from the 
Revivalists. 

4. The Oneida Communists, comprising those who believed in the neces¬ 
sity of human leadership as an auxiliary to direct divine guidance, were 
a selection from the Perfectionists. 

Some of the resulting characteristics of the Oneida Community group 
were the following: 

1. Hardiness. Until 1863 the Community members did the work of farm, 
factory and household with no hired help. They lived a pioneer life. The 


THE ONEIDA COMMUNITY EXPERIMENT 


377 


death-rate among them was only 72 per cent of that in New York State 
total population, 1910. (See table 1.) 

2. Longevity. Survival until eighty years of age and beyond was com¬ 
mon. Twenty-two died between the ages of eighty-five and ninety-six. 

3. Sufficient native ability to carry on successfully a diversified industry 
and the complex social affairs of a Community. 

4. Faculty of agreement. The members of the Community for forty 
years achieved the miracle of unanimity in all important moves. 

The 81 parents of the stirpicultural children may be classified oc¬ 
cupationally as follows: 

Men 


Founder and leader. 1 

Subordinate leaders. 4 

Superintendents of departments. 13 

Foremen. 3 

Mechanics, artisans and farmers. 13 

Salesmen, accountants and teachers. 5 

Inventor and dentist. 1 


Women 

Subordinate leaders. 

Superintendents of departments. 

Editors. 

Forewoman. 

Accountants, reporters and teachers. 

Compositors. 

Housewives. 


40 

2 

2 

3 

1 

7 

7 

19 

41 


It may be further mentioned that two of the superintendents were also 
editors, that several of those otherwise classified were important contrib¬ 
utors to the Community paper, and that eight of the forty men were 
college graduates. 

The standard of individual character aimed at was one in which first the 
spiritual, second the intellectual, third the moral, and fourth the physical 
departments of human nature were developed to the fullest extent com¬ 
patible with the above stated order of precedence. While the terms 
describing the four “departments of human nature” were not rigidly 
defined, yet they carry a meaning easily intelligible. Noyes’s own con¬ 
ception of them is stated in a “home-talk” dated April 5, 1869, as follows: 
















378 


H. H. AND G. W. NOYES 


Our nature is like a nest of four boxes. The inner one is the spiritual, the next is the 
intellectual, the next is the moral, and the outer one is the physical. God will begin to 
renew us by giving us ruggedness in the spirit, which will enable us to face all devils with¬ 
out fear and live in contact with them without contamination. Then he will give us 
ruggedness of the understanding, which fears no argument and overcomes all delusions. 
Then he will give us ruggedness of the moral nature, which keeps its resolutions and sub¬ 
mits to no condemnation. And finally through all these mediates he will give us rugged¬ 
ness of physical health. * 

The direction of the experiment during the early years was in the hands 
of an informal committee consisting of the so-called “central members” 
of the Community, Noyes himself exercising the preponderating influence. 
On January 25, 1875, a formal “Stirpicultural Committee” composed of 
six men and six women, two of the members being graduates of the Yale 
Medical School and the others persons of exceptional experience and judg¬ 
ment, was appointed by the Community and placed in charge. The 
“Stirpicultural Committee” functioned for about fifteen months, or until 
April 20,1876. After this the direction of the experiment passed again into 
the hands of the “central members” of the Community. 

The general methods of selection are indicated by the records of the 
“Stirpicultural Committee.” In a majority of cases application was made 
to the Committee by couples desiring to become parents, and the Com¬ 
mittee, after due consideration, would either approve or disapprove. If an 
application were disapproved, the Committee would always interest itself 
in an attempt to find a combination agreeable to those concerned which it 
could approve. Occasionally the Committee itself took the initiative in 
bringing about combinations which in their opinion were specially fit. 

The degree of selection exercised by the Committee may be judged from 
the following facts: 

1. The death-rate among the 81 selected parents was 22.5 per cent less 
than that of the group as a whole. (Compare tables 1 and 3.) 

2. The fathers of the stirpicultural children averaged 12.2 years older 
than the mothers. This indicated a certain amount of survival selection 
among the fathers, and accorded with the principle that quality of offspring 
is more dependent upon selection among fathers than among mothers. 

3. During a typical period of about fifteen months, out of 51 applications 
from men and women desiring to become parents, 9 were vetoed on the 
ground of unfitness, and 42 were approved. 

4. Exact statistics as to the number of combinations brought about by 
the initiative of the Committee are not available, but it is probable that 
about 25 per cent of the births were of this character. 


THE ONEIDA COMMUNITY EXPERIMENT 


379 


5. Although nearly as many fathers as mothers were engaged in the 
experiment, 30 of the fathers had only 1 child each, while the remaining 
10 had an average of 2.8 children each. 

Children were cared for by their mothers during infancy. When able 
to walk, a child was admitted to the day nursery department of the “ Chil¬ 
dren’s House,” the mother continuing the night care. From the beginning 
of the play stage until adolescence the “Children’s House” had complete 
charge, though parents visited their children and received visits from them. 
Much attention was given to diet, clothing, sanitation and profitable 
activity, and since epidemic diseases common in outside society were 
vigilantly excluded sickness among the children was rare. In case of 
sickness good medical attendance and the best of nursing were immediately 
available. There were also facilities for quarantine, night watchers, and 
appliances for comfort and convenience, such as few private families could 
afford. During the forty years 1841-1880 inclusive, in which a total of 
193 children were cared for, only 5 deaths occurred in the “Children’s 
House.” At adolescence a young person graduated from the “Children’s 
House,” and took his place in the general organization of the Community. 

At the re-organization of the Community as a joint-stock company on 
January 1, 1881, the provisions made for the women and children were 
these: 

The women of the Community participated equally with the men in the 
distribution of assets in the form of stock, the general basis of division being 
a proportionate part of the property brought in and an allowance for each 
year of membership. Several of the mothers who did not immediately 
marry were given a small extra cash allowance so long as they remained 
unmarried. 

Every child under two and a half years of age received an annual appro¬ 
priation of $200 in cash, until the age of two and a half was reached. 

Every child over two and a half and under sixteen years of age received 
an annual appropriation according to the profits of the Company, the 
minimum appropriation being $80 and the maximum $125 per year. 

Every child upon reaching its sixteenth birthday received $200 in cash. 

The Company obligated itself to maintain a suitable school for children 
under sixteen years of age free of charge. 

The Company also agreed to give preference to the members of the 
former Community and their children in the matter of employment. 

All these provisions were faithfully carried out by the new Company, 
and no cases of destitution among the women and children of the Com¬ 
munity have ever occurred. 


380 


H. H. AND G. W. NOYES 


A general survey of the Oneida Community stirpicultural experiment will 
show the following outstanding results: 

1. No mothers were lost during the experiment, nor indeed during the 
entire career of the Community, from causes directly or indirectly due to 
child-birth. 

2. No deaf and dumb, blind, crippled or idiotic children were ever born 
in the Community. 

3. In the group of 44 pre-stirpicultural children, born under the regime 
of birth-control but not of parent-selection, the death-rate was 82.6 per cent 
of that in the United States total population, 1870.* (See table 2.) 

4. In the group of 58 stirpicultural children born during the period 
1869-1879, 6 have died up to September 26, 1921. These 6 actual deaths 
represent 13.3 per cent of the expected deaths in the United States total 
population, 1870. (See table 4.) 

5. The causes of the 6 actual deaths in the group of stirpicultural children 
were these: 

Case 1. Age three days. Atelectasis after instrumental delivery. 

Case 2. Age seven years. Scarlet fever contracted during a residence in a large city. 

Case 3. Age nineteen years. Brain tumor. 

Case 4. Age thirty-nine years. Run over by a railway train. 

Case 5. Age forty-four years. Fall with aeroplane while flying for the allies in France. 

Case 6. Age forty-five years. Aortic Aneurism. 

6. Among the 52 survivors of the 58 stirpicultural births, now at ages 
forty-one to fifty-one there are two cases of sub-normal development. The 
first is a man fifty years of age, who has shown since birth a partial lack of 
muscular coordination, caused probably by cerebral hemorrhage during 
delivery. His mentality is normal, and his health, except for the defect 
noted, is good. He was for many years a foreman, and has always been 
able to support himself by his own labor. The second is a man forty-five 
years of age, whose mental development was retarded at about the age of 
seven by an accidental blow on the head which left a permanent indentation 
on the cranium. Though still slightly sub-normal mentally, he works hard on 
a farm and in a store, and enjoys reading. Physically he is well-developed. 
Aside from these two, there are no cases of even partial incapacitation for 
the normal activities of life. 

7. During the stirpicultural period a purposely increased birth-rate co¬ 
incided with a marked fall in the death-rate. (Compare tables 2 and 4.) 

8. In the group of 98 children, one or both of whose parents were stir¬ 
picultural, 5 actual deaths occurred during the period 1890-1920, 
representing 23.5 per cent of the expected deaths in New York State total 
population 1910. (See table 5.) 


THE ONEIDA COMMUNITY EXPERIMENT 


381 


9. In the group of 28 children, both of whose parents 1 were stirpicultural, 
no deaths have occurred during the period 1897-1920. In a group of 
equal size the number of expected deaths in New York State total population 
1910 is 6.2. (See table 6.) 

TABLE 1 


Comparison of actual and expected results among years of life exposed to risk, September 15, 
1880 , to close of 1920. 225 Oneida Community members sixteen years of age and 
above on September 15,1880. 


AGES ATTAINED 

YEARS OF 
LIFE 

EXPOSED 

ACTUAL 

DEATHS 

NUMBER OF DEATHS EXPECTED 

UNDER MORTALITY RATES OF FOLLOWING TABLES: 

United States 
Registration 
Area 

1910 

United States 
Registration 
Area Rural 
1910 

New York 
State total 
population 
1910 

United States 
population 
1870 
(Elliot) 

All ages 

5538.65 

166 

216.1 

188.5 

230.5 

227.2 

15 to 19 

43.00 

_ 

L 

0.1 

0.1 

0.1 

0.4 

20 to 24 

93.08 

1 

0.5 

0.5 

0.5 

1.1 

25 to 29 

150.00 

— 

0.9 

0.8 

0.9 

1.8 

30 to 34 

246.50 

3 

1.7 

1.4 

1.9 

3.2 

35 to 39 

349.00 

5 

3.0 

2.3 

3.4 

4.9 

40 to 44 

440.25 

1 

4.4 

3.2 

5.1 

6.9 

45 to 49 

540.16 

2 

6.8 

4.9 

7.8 

9.9 

50 to 54 

610.66 

5 

9.8 

7.0 

11.1 

13.5 

55 to 59 

634.98 

7 

14.8 

11.0 

16.6 

17.7 

60 to 64 

656.36 

14 

21.6 

16.7 

24.1 

23.9 

65 to 69 

603.41 

15 

28.6 

23.6 

30.9 

29.5 

70 to 74 

527.56 

31 

36.6 

32.5 

38.9 

35.5 

75 to 79 

344.99 

32 

35.1 

32.8 

35.9 

32.4 

80 to 84 

197.37 

23 

30.2 

29.6 

30.9 

26.2 

85 to 89 

87.33 

20 

18.1 

18.1 

18.5 

16.5 

90 and over 

14.00 

7 

3.9 

4.0 

3.9 

3.8 


Percentage, actual of expected mortality among 5,538.65 years of life exposed to risk. Sur¬ 
vivals of 225 Oneida Community Members sixteen years and above Jan. 1,1881, under 
observation from September 15, 1880, to close of 1920. 


TABLE USED AS “STANDARD" 

DEATHS 

EXPECTED 

PERCENTAGE 
ACTUAL DEATHS 
(166) OF 
EXPECTED 

United States total Registration Area, 1910. 

216.1 

76.8 

Rural part of United States Registration Area, 1910. 

188.5 

88.1 

New York State total population, 1910. 

230.5 

72.0 

Elliott United States, 1870. 

227.2 

73.1 


1 Complex marriage was abandoned August 28, 1879, since which date the members 
of the Community have married as in ordinary society. Of the 55 stirpicultural children 
who reached marriageable age, 18 intermarried, 32 married outside the group, and 5 
remained single. The results of the 50 marriages are shown in the text and in tables 
5 and 6. 































382 


H. H. AND G. W. NOYES 


TABLE 2 


Comparison of actual and expected number of deaths among survivors of 44 Oneida Commu¬ 
nity pre-stirpicultural births to Oneida Community pioneers. Period of observation , 

1847 to 1920 


AGE ATTAINED 

NUMBER 

OF YEARS 
OF LIFE 
EXPOSED TO 
RISK 

ACTUAL 

DEATHS 

NUMBER OF DEATHS EXPECTED 

ACCORDING TO DEATH RATES OF FOLLOWING TABLES: 

United 
States Reg¬ 
istration 
Area, 1910 

Rural part 
United 
States Reg¬ 
istration 
Area, 1910 

New York 
State 

total popu¬ 
lation, 1910 

United 
States total 
population, 
1870 
(Elliott) 

Massachu¬ 
setts Regis¬ 
tration 
Districts 
1855 
(Elliott) 

All ages 

1653 

27 

20.1 

16.0 

22.1 

32.7 

31.0 

Under 5 

179 

10 

6.5 

5.0 

6.9 

11.0 

12.5 

5 to 9 

166 

2 

0.6 

0.4 

0.6 

1.5 

1.3 

10 to 14 

163 

— 

0.4 

0.3 

0.4 

0.9 

0.7 

15 to 19 

159 

1 

0.6 

0.5 

0.6 

1.4 

1.4 

20 to 24 

140 

3 

0.7 

0.7 

0.7 

1.9 

1.7 

25 to 29 

140 

1 

0.5 

0.7 

0.9 

1.7 

1.7 

30 to 34 

134 

2 

1.0 

0.8 

1.0 

1.7 

1.6 

35 to 39 

124 

3 

1.1 

0.8 

1.2 

1.7 

1.6 

40 to 44 

108 

1 

1.1 

0.8 

1.3 

1.7 

1.5 

45 to 49 

98 

2 

1.2 

0.9 

1.4 

1.8 

1.5 

50 to 54 

94 

— 

1.5 

1.1 

1.7 

2.1 

1.2 

55 to 59 

72 

1 

1.7 

1.3 

1.9 

2.0 

1.6 

60 to 64 

42 

— 

1.4 

1.1 

1.5 

1.5 

1.3 

65 to 69 

25 

1 

1.2 

1.0 

1.3 

1.2 

1.0 

70 to 74 

9 

— 

0.6 

0.6 

0.7 

0.6 

0.4 


Percentage , actual of expected number of deaths. Actual deaths = 27 


MORTALITY TABLE USED AS “STANDARD” 

NUMBER 

OF DEATHS 
EXPECTED 

PERCENTAGE 

ACTUAL 

OF EXPECTED 
DEATHS 

United States Registration Area, 1910. 

20.1 

134.3 

Rural part, United States Registration Area, 1910. 

16.0 

168.8 

New York State population, 1910. 

22.1 

138.1 

United States total population, 1870. 

32.7 

82.6 

Massachusetts Registration Districts, 1855. 

31.0 

87.1 
































THE ONEIDA COMMUNITY EXPERIMENT 


383 


TABLE 3 


Comparison of actual and expected number of deaths among survivors of 79 of the 81 parents 
of stirpicultural children. Births , 1809 to 1858. Observations carried to 

August 16, 1921 



NUMBER OF 


NUMBER OF DEATHS EXPECTED: 

AGES ATTAINED 

YEARS OF LIFE EX¬ 
POSED TO RISK 

ACTUAL DEATHS 

New York State 
total population, 
1910 

United States 
population, 1870 
(Elliott) 

All ages, 20 to 89 

3931 

52 

.92.9 

102.7 

20 to 24 

395 

_ 

2.0 

4.6 

25 to 29 

391 

1 

2.4 

4.8 

30 to 34 

390 

1 

3.0 

5.0 

35 to 39 

381 

2 

3.7 

5.3 

40 to 44 

375 

— 

4.3 

5.9 

45 to 49 

372 

2 

5.4 

6.8 

50 to 54 

360 

2 

6.6 

8.0 

55 to 59 

348 

4 

9.1 

9.7 

60 to 64 

320 

7 

11.8 

11.7 

65 to 69 

283 

8 

14.5 

13.9 

70 to 74 

189 

13 

13.9 

12.7 

75 to 79 

83 

7 

8.6 

7.8 

80 to 84 

30 

2 

4.7 

4.0 

85 to 89 

14 

3 

3.0 

2.6 


Percentage, actual of expected number of deaths. Actual deaths = 52 


MORTALITY TABLE USED AS “STANDARD” 

NUMBER 

OF DEATHS 
EXPECTED 

PERCENTAGE 

ACTUAL 

OF EXPECTED 
DEATHS 

New York State total population, 1910. 

92.9 

56.0 

United States total population, 1870. 

102.7 

50.6 






















384 


II. H. AND G. W. NOYES 


TABLE 4 


Comparison of actual and expected number of deaths among survivors of 58 stirpicultural 
births in the Oneida Community. Births, 1869 to 1879. 

Observations carried to September 26, 1921 


AGES ATTAINED 

NUMBER 

OF YEARS 

OF LIFE 
EXPOSED TO 
RISK 

ACTUAL 

DEATHS 

NUMBER OF DEATHS EXPECTED 

UNDER MORTALITY RATES OF SPECIFIED TABLE: 

Rural part 
of United 
States regis¬ 
tration area, 
1910 

United States 
Registration 
Area total 
table, 1910 

New York 
State total 
population, 
1910 

United States 
total popu¬ 
lation, 1870 
(Elliott) 

All ages 

2638 

6 

19.7 

24.7 

26.6 

45.2 

Under 5 

285 

1 

18.0 

10.3 

11.0 

17.7 

5 to 9 

283 

1 

0.7 

0.9 

1.0 

2.6 

10 to 14 

280 

— 

0.6 

0.6 

0.6 

1.6 

15 to 19 

280 

— 

0.9 

1.0 

1.0 

2.4 

20 to 24 

275 

1 

1.4 

1.4 

1.4 

3.2 

25 to 29 

275 

— 

1.4 

1.6 

1.7 

3.4 

30 to 34 

275 

—. 

1.6 

1.9 

2.1 

3.5 

35 to 39 

275 

— 

1.8 

2.4 

2.6 

3.8 

40 to 44 

255 

1 

1.9 

2.6 

2.9 

4.0 

45 to 49 

134 

2 

1.2 

1.7 

1.9 

2.5 

50 to 54 

21 

— 

0.2 

0.3 

0.4 

0.5 


Percentage , actual of expected number of deaths among survivors of 58 stirpicultural births 
in the Oneida Community. Expected deaths computed from mortality 
rates of specified tables. Actual deaths = 6 


MORTALITY TABLE USED AS “STANDARD” 

NUMBER 

OF EXPECTED 
DEATHS 

PERCENTAGE, 
ACTUAL (6) 

OF EXPECTED 
NUMBER 

OF DEATHS 

Rural part of United States Registration area, 1910. 

19.7 

30.5 

United States Registration Area, 1910. 

24.7 

24.3 

New York State total population, 1910. 

26.6 

22.6 

United States total population, 1870 (Elliott). 

45.2 

13.3 




























THE ONEIDA COMMUNITY EXPERIMENT 


385 


TABLE 5 


Comparison of actual and expected number of deaths among survivors of 97 births, 
one or both of whose parents were stirpicultural. Births, 1890 to 1917. 
Observations carried to 1921. 


AGES ATTAINED 

NUMBER OF YEARS 

OF LIFE EXPOSED TO 
RISK 

ACTUAL DEATHS 

NUMBER OF DEATHS 
EXPECTED 

New York State total 
population, 1910 

All ages 

1480 

5 

21.3 

Under 5 

464 

4 

18.0 

5 to 9 

428 

— 

1.5 

10 to 14 

345 

— 

0.8 

15 to 19 

170 

— 

0.6 

20 to 24 

61 

1 

0.3 

25 to 30 

12 

— 

0.1 


Percentage, actual of expected number of deaths 
New York State population, 1910 as “standard”: 23.5 

TABLE 6 


Comparison of actual and expected number of deaths among survivors of 28 births, both of 
whose parents were stirpicultural. Births, 1897 to 1917. Observations carried to 1921. 



NUMBER OF YEARS 


NUMBER OF DEATHS EXPECTED 

AGES ATTAINED 

OF LIFE EXPOSED 

TO RISK 

ACTUAL DEATHS 

New York State 
total population, 
1910 

United States 
population, 1870 
(Elliott) 

All ages 

412 


6.2 

10.7 

Under 5 

138 

_ 

5.3 

8.5 

5 to 9 

124 

— 

0.4 

1.2 

10 to 14 

96 

— 

0.2 

0.5 

15 to 19 

42 

— 

0.2 

0.4 

20 to 24 

12 

-■ 

0.1 

0.1 


Percentage, actual of expected number of deaths. Actual deaths = 0 


MORTALITY TABLE USED AS “STANDARD” 

NUMBER OF DEATHS 
EXPECTED 

PERCENTAGE, 
ACTUAL OF 
EXPECTED DEATHS 

New York State total population, 1910. 

6.2 

None to be 

United States total population, 1870. 

10.7 

computed 

































TABLE 7* 


Summary: Comparison of actual and expected number of deaths in various Oneida 

Community groups 


GROUP 

ACTUAL 

DEATHS 

EXPECTED 
DEATHS 
ACCORDING 
TO CON¬ 
TEMPORARY. 
TABLES 

PERCENT¬ 
AGE, ACTUAL 
OF 

EXPECTED 

NUMBER 

OF DEATHS 

44 pre-stirpicultural births. 

27 

32.7 

82.6 

225 Oneida Community members, 16 years of age and 
over on September 15, 1880. 

166 

227.2 

73.1 

Parents of the stirpicultural children. 

52 

102.7 

50.6 

Children, one or both of whose parents were stirpicultural 

5 

21.2 

23.6 

The stirpicultural children. 

6 

45.2 

13.3 

Children both of whose parents were stirpicultural. 

— 

6.2 

— 


* The Scandinavian method of computing the ratio of actual to expected mortality 
has been used in these tables rather than the method of computing the death rates per 
thousand as being more accurate for small groups. The death rates per thousand used in 
computing the expected deaths are given below: 

Death rates per 1000 at specified central ages of five year age groups. Population mortality 
tables used to test Oneida Community mortality data 


DEATH RATES PER 1,000 


AGE GROUP 

United States 
Registration 
Area, 1910 

Rural part of 
United States 
Registration 
Area,1910 

New York 
State, total 
population, 
1910 

United States 
total popu¬ 
lation, 1870 
(Elliott) 

Massachusetts 
Registration 
Districts, 1855 
(Elliott) 

Under 5 

36.29 

27.96 

38.72 

61.92 

69.84 

5 to 9 

3.30 

2.51 

3.55 

9.27 

7.78 

10 to 14 

2.22 

2.02 

2.17 

5.66 

4.35 

15 to 19 

3.52 

3.29 

3.52 

8.50 

8.78 

20 to 24 

5.19 

5.13 

5.10 

11.75 

10.75 

25 to 29 

5.85 

5.24 

6.10 

12.18 

11.88 

30 to 34 

7.09 

5.66 

7.71 

12.87 

12.19 

35 to 39 

8.59 

6.48 

9.62 

13.97 

12.58 

40 to 44 

10.09 

7.26 

11.53 

15.67 

13.43 

45 to 49 

12.63 

9.02 

14.41 

18.25 

14.81 

50 to 54 

16.01 

11.54 

18.21 

22.12 

18.36 

55 to 59 

23.37 

17.26 

26.15 

27.89 

22.32 

60 to 64 

32.96 

25.37 

36.73 

36.42 

31.94 

65 to 69 

47.41 

39.04 

51.14 

48.96 

39.73 

70 to 74 

69.38 

61.56 

73.70 

67.30 

48.38 

75 to 79 

101.74 

94.95 

104.14 

94.01 

107.05 

80 to 84 

153.06 

150.08 

156.72 

132.77 

166.26 

85 to 89 

206.84 

206.71 

211.35 

188.82 

229.82 

90 to 94 

279.90 

285.86 

276.04 

269.69 

280.34 

95 to 99 

354.55 

382.71 

365.31 

386.03 

315.78 


We wish to express our hearty appreciation for the assistance on the construction of 
the mortality tables given by Mr. Edwin W. Kopf, assistant statistician of the Metropoli¬ 
tan Life Insurance Company. 


386 






























SOME FAMILIES AS FACTORS IN ANTI-SOCIAL CONDITIONS 
AMOS W. BUTLER 

Indiana Board of State Charities, Indianapolis 

One of the first undertakings of the Board of State Charities of Indiana, 
after it was established in 1889, was to ascertain how many persons were 
inmates of the state and local institutions and, later, who these persons were, 
some facts of personal history and the reason they were public charges. The 
information, collected by means of institution reports, was transferred to 
a card registration. Beginning modestly with the inmates of ninety-two 
county poor asylums and four state hospitals for the insane, the registra¬ 
tion contained abut 5700 names at the end of its first year. From time to 
time other institutions were added, and the reports were made to include 
something of both personal and family history. All this was properly 
carded. 

The registration now contains over 158,000 names of persons who are, 
or within the past thirty-one years have been, inmates of eighteen state 
charitable and correctional institutions, ninety-two county poor asylums 
and thirty-three orphans’ homes. It is maintained in duplicate, one set 
being arranged by institutions, the other alphabetically and phonetically. 
It is the latter that brings family names together, and no one can glance 
over the cards without being impressed by the frequency with which feeble¬ 
mindedness or a related defect appears on these records. It was from them 
in 1896 that Mr. Ernest P. Bicknell, then secretary of the board, made a 
study of 248 families, selected because of feeble-mindedness in one or more 
generations. Concerning these records, Mr. Bicknell well said: 

They are not clean cut, not properly rounded out. They begin in obscurity, come into 
view for a few years, and fall back into obscurity again. But the broken stories of their 
misery, their perpetuation of their own wretched kind, their demoralizing influence upon 
their fellows, their dragging down of the average of morality, intelligence, and physical 
development, are sorrowful beyond words. 1 

Eleven years afterward it was my privilege to present to the National 
Conference of Charities and Correction a similar study from this registra- 

1 National Conference on Charities and Correction, 1896, page 219. 

387 


388 


AMOS W. BUTLER 


tion, of 803 families, more than half the members of which were 
feeble-minded. 2 

Beside this card registration of institution inmates, the board has a 
record, by name, of every family aided by official out door poor relief, of 
every person received at a county jail, of every child born in a maternity 
hospital. All these make available a vast amount of information concern¬ 
ing public dependents. Out of it has grown a desire for still more data, 
and one result was the appointment by the governor in 1915 of a com¬ 
mittee to study the whole problem of mental defectives and suggest a 
program for their care. 

This committee has surveyed ten of our ninety-two counties, listing every 
feeble-minded, epileptic or insane person, found 5322 in all, an average of 
2.1 per cent of the population of those ten counties. On this basis, Indiana 
has 56,718 mental defectives—44,284 feeble-minded, 8311 insane and4123 
epileptics. It is estimated that 25,232 of these do not need institutional 
care, but of those who do need it, the state is now caring for 79 per cent 
of the insane, 23 per cent of the epileptics and but 7 per cent of the feeble¬ 
minded. It is plain, therefore, that it is the feeble-minded who constitute 
our most serious problem. We have but two state institutions for them a 
school at Fort Wayne, a colony farm at Butlerville. Together they have 
about 1500 inmates—a very small porportion of the number in our state 
who need institutional care. Where are the rest? They are in institutions 
not suitable for them, or they are at large in the communities. 

More than half the inmates of our county poor asylums are mental 
defectives. They wander in and out almost at will. Usually present in 
large numbers in the winter months, they leave when spring comes, and 
roam about the country, satisfied with whatever offers shelter,—an old hut 
or sometimes a rail pen. 

Because the state school for feeble-minded youth is crowded, our orphans’ 
homes are gradually filling up with children who cannot be placed with 
foster parents. A study of 225 dependent children in one institution 
disclosed 22.6 per cent mental defectives. 

Many of our prisoners are feeble-minded. “ Mental defect is good soil 
in which to develop crime.” The ten counties surveyed by our com¬ 
mittee were represented in the Indiana State Prison by 69 men, of whom 
60.8 per cent were mental defectives. 

Of nearly 1000 school children tested by the Binet-Simon method, from 
2 to 3 per cent were found feeble-minded and 10 per cent in need of special 

2 N.C.C.C., 1907, page 1. 


FAMILIES AS FACTORS IN ANTI-SOCIAL CONDITIONS 


389 


instruction. They were receiving no benefit themselves in the public 
schools, and were hindering the progress of normal pupils. 

In our records of official outdoor relief, we usually expect to find defectives 
in about one-fourth of the twenty thousand families aided annually. 
These people are living in their own homes, or in some deserted shanty, 
depending on charity, to eke out a miserable existence. 

What kind of homes do they have? Let me quote from an investigator’s 
report on some of them: 

The dirt, disorder, filth and cluttering are indescribable. Animals wander in and 
out. Garbage on the table, piles of filthy clothing on the floor, food stored under beds, 

an awful stench arising from the filth.The yard of one house a jungle of 

harness, ropes, old buggies, broken cots, cans, old iron, wheels. In the house the beds 
are piled high with dirty rags. One room so cluttered with old furniture that a path¬ 
way to the beds is the only unoccupied space.Many homes dilapidated. 

Isolation remarkable. Roads are just ruts and ditches. 

The records which have been accumulating in our office for the past 
quarter of a century, and more recently those collected by the Committee 
on Mental Defectives, while somewhat in the nature of a census, yet contain 
in some cases many generations of family history. Now and then when 
a particular name occurred with unusual frequency or a particular locality 
seemed unduly represented in public institutions, a special investigation 
has been made. In one way and another, hundreds of degenerate families 
have been listed, some going back as far as seven generations. Among 
them are scores of references to the Tribe of Ishmael, which is to Indiana 
and the central west what the Jukes are to New York and the Kallikaks to 
New Jersey, though a much larger group than these or any other that has 
so far been studied. While these family records are strikingly similar, some 
are noted for a particular trait or tendency. There is, for example, a family 
in eastern Indiana, of which the mother and ten of her eleven children are 
mentally unbalanced. Two sons and two daughters of this family com¬ 
mitted suicide. Another daughter with two children threatened their lives 
and her own. Their maternal grandfather met death by his own hand. 

A family which has long been a veritable hotbed of immorality lives in 
southern Indiana. One woman, whom we will call “Polly,” is known to have 
eleven illegitimate children, each with a different father. One of Polly’s 
daughters, feeble-minded like her mother, has had eight illegitimate children, 
seven of whom are of the same mental caliber. One of these seven has had 
four illegitimate children. In this one group there have been twenty-three 
illegitimate children, the offspring of three feeble-minded women. Al¬ 
together, Polly has fifty-six lineal descendants, thirty-one of whom are feeble- 




390 


AMOS W. BUTLER 


minded and eighteen of whom have been inmates of public institutions. 
Sixteen of the eighteen are known to have spent a total of seventy-two years 
on public support, at a cost of $10,800. This is one branch of a family 
group of 477 individuals representing seven generations. The younger 
members are still a serious problem in the life of their community. 

In another county is a family group of 152 individuals, notorious not only 
for shiftlessness and immorality, which so often characterize these records, 
but also for crime. Fourteen of its members have police court records, 
three have been convicted for serious crimes, one has been in a reform 
school. 

The writer once had ranged before him in a county poor asylum four 
feeble-minded persons: a man, his daughter, grand-daughter and great 
grandson—four generations, all living at public expense. 

Thirty years ago a young feeble-minded woman was admitted to one of 
our county poor asylums. A year later a blind man became an inmate. 
The two became acquainted and in time were married. Their wedding 
occurred in the poor asylum, they set up their family altar there and to 
them were born five children, all in the institution. Quite recently one of 
their daughters was committed to the Woman’s Prison for perjury, in 
connection with statements concerning her illegitimate child, who is now 
in an orphan’s home. In this case, as not infrequently happens, public 
officials connived at that which later brought a heavy burden of expense 
and disgrace upon their community. They “ knew not, and knew not that 
they knew not. ” 

One could fill a book with these Indiana stories of misery and degradation, 
of sin and suffering and crime, of public ignorance, indifference and neglect. 
And they could doubtless be duplicated in ever state in the Union. Every¬ 
where our people have failed to realize what was happening and these 
weaker children of the land have grown incredibly strong in numbers and 
in power for evil. More than once I have taken occasion to say that feeble¬ 
mindedness is one of the most potential destructive factors in our civiliza¬ 
tion. It produces more pauperism, more crime, more degeneracy, than 
any other one force. It is a fact we have to face, a condition we have to 
meet, a power we must keep under. 


PEDIGREES OF PAUPER STOCKS 


E. L. LIDBETTER 
London, England 

In asking me to prepare a paper on this subject for the Second Interna¬ 
tional Eugenics Congress, Major Leonard Darwin no doubt had in view the 
fact that I have been engaged in an investigation as to family pauperism 
for many years past. 

As a result a considerable number of pedigrees—some 400 to 500—have 
either been prepared or are in course of preparation. The examples here 
submitted are the more interesting of those already completed. 

As the Congress is of an international character it is necessary to say 
that the term “pauperism” is used in this paper in its strictly legal and offi¬ 
cial sense as understood in England. That is to say only such persons are so 
described as are or have been maintained wholly or in part by funds raised 
out of public rates levied and administered by a board of guardians, duly 
elected for that purpose by the ratepayers. All other forms of public and 
private assistance are disregarded in the pedigrees. 

Until a few years ago it would have been possible to say that all assist¬ 
ance afforded out of public funds was administeredby the poor law guard¬ 
ians and therefore fell within the term “pauperism.” This, however, is 
no longer the case, vast sums of public money being now expended by 
public authorities other than the poor law guardians, among which may be 
mentioned: 

a. Old age pensions, administered by special commissioners. 

b. Feeding of school children by education authorities. 

c. Relief afforded by public health authorities in cases of sickness and 
maternity. 

d. Maternity homes and other non-contributary hospitals provided by 
public health authorities out of local rates. 

e. Unemployment relief provided from imperial funds (i.e., from taxa¬ 
tion and not from local rating). The latter nominally a contributary 
scheme of unemployed insurance, but actually as now administered non- 
contributary relief in unemployment—in many cases supplemented by the 
Poor Law Guardians out of local rates. 

391 


392 


E. L. LIDBETTER 


To those who have not given careful attention to the subject the prepara¬ 
tion of pedigrees in relation to pauperism seems to imply that pauperism as 
such is hereditary. No such proposition is put forward in this paper. But 
it is contended that the pauperism is a consequence of inherent and trans¬ 
missible defects of character, and that pauperism is dependant upon mental, 
moral and physical defects, standing in relation to the pauperism as cause to 
effect, such as render the individual unable to hold his own in the competi¬ 
tion for existence so that he fails to be at all times self-supporting—often 
for considerable periods. 

To a gathering assembled in the name of Eugenics it will not be necessary 
to argue at length that pauperism is to be attributed to bad heredity, either 
in the sense of the actual transmission of defect, or the perpetuation of low- 
grade types. The really important question is as to what proportion of 
existing pauperism can be shown to be due to those causes. In this un¬ 
fortunately there exists no data for our guidance, and I regret that I am 
unable to supply that deficiency. 

In 1913 I commenced an investigation which would, if completed, have 
been of assistance in this respect; but the enquiry was unfortunately aban¬ 
doned when war conditions made it impossible to carry it on. In passing, 
it should be observed that the gradual disintegration of the Poor Law service 
now going on by the introduction of parallel services as indicated above, 
will greatly hamper—if not render impossible—any attempt at a comprehen¬ 
sive investigation in the future. The complete coordination of the whole 
public assistance service is essential to such an investigation. 

In these circumstances it must be understood that the diagrams submitted 
herewith of particular cases are selective. They are unquestionably repre¬ 
sentative of a great mass of pauperism in England—particularly in the great 
towns. But it is not possible to say what proportion of the whole they 
represent. 

In all the official investigations with regard to pauperism in this country 
no attempt has ever been made to analyse or define the personality of the 
pauper, his character, antecedents or capacity. Mass statistics have been 
produced, reduced to some rough classification and inferences drawn from 
the result. The effect has been to obscure the essentially personal element, 
so that the problems involved have come to be regarded as purely economic 
problems; and no doubt much modern legislation is based upon this wholly 
mistaken view. 

All engaged in the administration of the Poor Laws have long been struck 
by the tendency of particular individuals and particular families to apply 
to the relieving officer in almost every domestic contingency, and indeed 


PEDIGREES OF PAUPER STOCKS 


393 


for whole families to spend their lives in intermittent chargeability. To 
what extent does this practice obtain and in what proportion does it stand 
to the sum total of pauperism? 

Taking the inmates of the workhouse and hospital of a small parish in the 
east end of London on a particular day in the year 1913 the following figures 
show the number of those present on that day and the number of times 
which the several individuals had been previously admitted as inmates: 


NUMBER OF TIMES ADMITTED 

NUMBER IN 
WORKHOUSE 

NUMBER IN 
INFIRMARY 

TOTAL 

First admission. 

445 

240 

685 

Second admission. 

197 

74 

271 

Third admission. 

106 

41 

147 

Fourth admission. 

62 

25 

87 

Fifth admission. 

39 

18 

57 

Sixth admission. 

29 

9 

38 

Seventh admission. 

11 

7 

18 

Eighth admission . 

18 

7 

25 

Ninth admission. 

11 

3 

14 

Tenth admission. 

11 

2 

13 

More than 10 and under 20. 

26 

13 

39 

More than 20 and under 30. 

16 

5 

21 

More than 30 and under 50. 

11 

1 

12 






Among workhouse cases, 1 each of 53d, 63d, 77tfi, 96th, 110th, 113th, 
130th, 152d, and 184th time of admission. Among infirmary cases, 1 each 
of 55th and 115th time of admission. These latter would of course be 
extraordinary cases. Mainly men allowed out from day to day to look for 
work—who seldom found it. 

I would illustrate this latter point by giving the details of a particular 
case: 

Case of man Owen. Born 1859. Single. Cabinet maker. First entered workhouse 
in February 1905, at age of forty-six years. Was “in and out” on 108 occasions and 
finally left in January, 1916, when the war-time call for men had reduced the standard of 
employable labor. During this time the longest period out of the workhouse was 271 
days and the longest period in as an inmate was 231 days. In the whole period there were 
3990 days: he spent 2397 of these in the house and 1593 out. Average duration of each 
stay in the workhouse a fraction over 22 days, average stay out of it a fraction over 15 
days. 

On 106 occasions (not counted in the above figures) he discharged himself in the 
morning to look for work and returned the same evening. If these are added to the 
other 108 admissions it gives 214 occasions upon which the official machinery attending 
admission to the workhouse was put into operation by this man in a period of 10 years and 
11 months, out of which he spent a total of 6 years, 6 months and 20 days in the work- 
house. 






















394 


E. L. LIDBETTER 


The generality of these figures (p. 393) would no doubt have been greatly 
reduced by an examination of the family relations of the individuals 
concerned. This was in fact attempted, but the variety of the relationship 
and the labour required for the investigation was so considerable that the 
attempt broke down and had to be abandoned—one of the consequences of 
the outbreak of war. 

The symbols used in the preparation of these pedigrees are as follows: 

© o o® o 

4 5 6 7 8 

Letters added to the symbols: 

T = Tuberculous. 

B = Blind. 

E = Epileptic. 

D = Deaf and dumb. 

6. Insane pauper. 

7. Died in infancy. 

8. Imbecile. 

Note. Where the lines denoting relationship are disconnected or broken (thus, 
-)it indicates that the parents were not married. 

Although the actual pedigree charts are not reproduced in this paper some idea of their 
findings can be gathered by an examination of the pedigree-summaries of the following 
families: 

Family I. A Group of Mildly Incompetent Persons in Six Generations. 

Very Little Sickness Indicated 


Paupers: 


Non-paupers or unknown: 


Chronic. 

.. 13 

Died in infancy. 

...21 

Occasional. 

.. 64 

Known non-paupers. 

...55 

Insane. 

.. 6 

Unknown. 

...31 

Reared in poor law schools. . 

.. 7 



Total. 

.. 90 

Tuberculous. 

...10 



Epileptic. 

... 3 



Blind. 

... 1 


On the date when the investigation commenced in 1913, seven of these were charge¬ 
able: six in the area under investigation and one beyond. These seven have a col¬ 
lective period of chargeability amounting to 66 years and seven months. Others in 
the group between them (as far as can be ascertained) 142 years; giving a total in the 
group of 275 years. 

The four brothers shown in the third generation at 8, 9, 10, and 11 are representative 
of the whole. 

No. 8. Entered workhouse at 60 years of age: was out for two short periods, average 
35 days each, and had at the date of enquiry been in the house continuously for six 
years and 5 months. Total chargeable period at that time 7 years and 11 months. 


O 

1 2 3 

1. Normal. 

2. Particulars not known. 

3. Occasional pauper. 

4. Chronic pauper. 

5. Reared in pauper schools. 














PEDIGREES OF PAUPER STOCKS 


395 


No. 9. Entered house at 43 years of age, and has not since been out. Chargeable 
period 26 years. Has a club foot. 

No. 10. Entered at the age of 34 years. Total period chargeable on the appointed 
day 15 years and 8 months. Last period unbroken 7 years and 7 months. Has been 
out 22 times, average stay out 7 months and 1 day. Total period out since he first came in 
13 years 2 months 13 days. 

No. 11. Entered at 26 years of age. Total period chargeable at the appointed day 
20 years, 9 months, 17 days. Last period unbroken 4 years. Has been out in all 60 
times, average 2 months 13 days. Aggregate period out since he first came in 12 years, 3 
months 13 days. 

Of these four brothers two were unmarried, and the others had wives and children. 
The wives carried on the homes after the men entered the workhouse. 

Family II. A Group of Vicious Non-moral Individuals 

Illegitimacy. Crime and Desertion. This illustrated by the large number of children 


reared in Poor Law schools. 

Paupers: 

Chronic.20 

Occasional. 44 

Insane. 3 

Reared in Poor Law schools. 26 

Total.93 


Family III. Insanity and Eye Disease in Six Generations 


Paupers: 


Chronic. 

. 2 

Non-paupers: 


Occasional. 

...... 56 

Normal. 

.30 

Blind. 

. 7 

Not known. 

.23 

Insane. 

. 10 

Died in infancy. 

. 10 

Blind and insane. 

. 5 




— Not known includes 13 parents of 81 
Total.80 children. 


Professional men and others interested in Family III will find it set out in 
greater detail in “ Brain” volume 35, part 3 (1913). 

These three large families have been selected for this paper because they 
show in such a clear way the three distinct classifications in these groups— 
the inheritance of mental disorder, the low grade type of mildly incompe¬ 
tent persons, and the distinctly non-moral group. Generally speaking these 
classes are found in combination all in the same group, but here they are 
found clearly defined and are so presented. 

From this point of view it is particularly interesting to notice their com¬ 
bination in marriage and reproduction, for not only do they reproduce as 
the biological students would expect, but like attracts like in marriage to 
















396 


E. L. LIDBETTER 


a remarkable degree. It needs only a paternal Poor Law system, easy of 
access, to ensure the survival value of these types and their multiplication. 

Only a want of space and time prevent me showing other pedigrees as 
large and important as these. I would point out too that these are pedigrees 
collected in the area of a single Poor Law authority—and that a very small 
one. The pauperism shown is of course the minimum. In few parishes 
are records available for more than thirty years and in fewer still are they 
intelligible over that period. It follows that there may have been in each 
pedigree a larger number of paupers than is here shown. Furthermore the 
period of the earlier generations shown is back to the limit of the memory 
of the available witnesses—another limiting difficulty. 

It is necessary to define more clearly the attitude of Eugenics to pauperism. 
In so far as pauperism is of a temporary character—as for instance in non¬ 
recurrent temporary sickness, or relief for the family of a man bona fide 
unemployed, or in the case of orphans maintained in Poor Law Schools 
whose parents were not suffering from diseases known to be hereditary—we 
are concerned in the matter only to the extent that eugenics requires the 
best environment for the biologically desirable. This is, however, a sort of 
side issue of eugenics proper and many no doubt will deny that it has any 
relation to eugenics. We are mainly concerned with the poor law in so far 
as pauperism is in relation to hereditary defect and its reproduction. Two 
questions are cardinal to the issue before us. Can it be shown that there is 
in existence a definite type of pauper family? Secondly and if so, is the 
pauperism in those families associated with a definitely transmissible charac¬ 
teristic standing in relation to the pauperism as cause and effect? 

I think there can be no doubt that the figures and the pedigree-summaries 
I have put before you answer, and abundantly answer, the first question 
in the affirmative. 

With regard to the second question the pedigree-summaries have been 
specially selected because they do show in each case a definitely transmissi¬ 
ble characteristic not only associated with pauperism, but standing in rela¬ 
tion to it as cause and effect. It is of the very greatest importance in this 
investigation that we should realize that whenever these pedigrees have been 
developed to a sufficient size, in each case the distinctive quality of that case 
has become apparent. This is the most important fact brought out by this 
series of cases. Not only is the distinguishing characteristic observable in 
each generation, but the inter-marriages are with the same type until the 
pedigree broadens out and is absorbed in the general community. 

In addition to these three large pedigree-summaries, I have described 
and charted four smaller ones, which should be of some interest to Mende- 
lians and others interested in indirect transmission. 


PEDIGREES OF PAUPER STOCKS 


397 


I would summarise this paper by submitting the following general 
propositions: 

1. That there is in existence a definite race of chronic pauper stocks, 
intermingled with the general community, not recruited to any large extent 
from the normal population, and not sensibly decreased by the agencies for 
the promotion of human efficiency. (The proportion this class bears to the 
total number of persons chargeable is unknown, but it must be considerable 
and some enquiry should be instituted to determine the fact). 

2. That modern methods of public and private charity tend to encourage 
the increase of this class by relieving the parents of the normal responsibili¬ 
ties of parenthood. 

3. That the tendency of all modern movements in matters relating to the 
care and upbringing of children, is and must continue to be towards a 
very large increase in parental responsibility, and as this movement goes on, 
the right of the community to decide as to the kind of children it is to shelter 
and maintain should increase also. 

4. That the development of Poor Law administration in the future should 
be towards its perfection as an organisation for the reception and care of 
those mental and physical degenerates, whom it now seems that all human 
communities must produce. 

5. That the reduction of this class may be brought about by a due ob¬ 
servance of the laws of heredity, so far as they are surely known, and that 
that reduction may become progressive in proportion as our knowledge 
grows. 


THE TRIBE OF ISHMAEL 


ARTHUR H. ESTABROOK 

Eugenics Record Office, Carnegie Institution of Washington 

In 1877 Dugdale published the story of the Jukes, calling it a study on 
Crime, Pauperism, Disease and Heredity, and giving in it the sociological 
history of six hundred people, all descended from five sisters, showing 
how the same anti-social traits have continually appeared for six generations. 
He said that these “ Jukes” were but a few of a big class. 

A little later, in the same year, a minister of the gospel in Indianapolis, 
Indiana, read the Juke book and was very much impressed by the descrip¬ 
tion of the folk and the generalization drawn by Dugdale. At about the 
same time he found the following case of destitution in that city. The 
following is quoted from his diary: 

A family, composed of a man, two children, the woman’s sister and child, the man’s 
mother, blind, all in one room ten feet square. One bed, a stove, no other furniture. 
When found they had no coal, no food. Dirty, filthy, because of no fire, no soap, no towels. 
It was the most abject poverty. I have never seen anything like it. We carried supplies 
to them. 

Two days later in his diary: 

The case alluded to under the date of Friday seems to be a case similar to that of 
the “Jukes.” I went to the office of the township trustee and found them under the 
name of the Ishmaelites. 

The story of how the name “ Ishmaelite” had come on the books of the 
township trustee is of interest. I may add at this point that in Indiana 
the township trustee is the official whose duty it is to oversee the giving of 
poor relief. From 1800 to 1850 many families from eastern Kentucky 
and southern Ohio for divers reasons were moving west. Many of these 
were the industrious, ambitious, hard-working peoples who were moving 
on to make a new home “further west.” These are not of interest to us 
sociologically. Some of these had left their homes with no preparation for a 
journey across what was in those days almost a wilderness. And so many of 
them were forced to beg for food and sustenance when they came to a settle¬ 
ment. Indianapolis was the first big “stopping place,” and the people 

398 


THE TRIBE OF ISHMAEL 


399 


were generous. And so Indianapolis came to be noted as a place where the 
people were friendly and very willing to help others who too were moving 
west into new habitats. This friendliness and generosity were soon to be 
taken advantage of and wanderers came to Indianapolis knowing that they 
would be cared for. During this period then Indianapolis became the home 
of many families who were dissatisfied with the harder struggle for existence 
in the developing country in Kentucky and Ohio and those who had the 
restless wandering spirit. These latter could travel when the weather was 
good and return to Indianapolis for the winter and live through this period 
by begging in the city just as in the summer they begged their way on the 
roads. There were quite a number of these families who disregarded the 
law of the land, who did much as they wanted to do and cared little for the 
opinion of others. They wandered from place to place. The men were 
shiftless; the women, immoral, and the children, ill-fed and clothed, the typi¬ 
cal feeble-minded people who are so easily recognized today. How many 
of these families there were is not known but a rough estimate would place 
it at not less than four hundred. It is these families and their offspring 
and interconnections that formed the greater part of the pauper population 
of Indianapolis of years ago. 

The amount of poor relief given in Indianapolis had become so great that 
in 1876 a systematic effort was made to study this pauperism in the city and 
make plans to reduce it. A system of historical records was devised cover¬ 
ing the names and social condition of these applicants for relief; making 
special note of all relatives, this for the purpose of finding any who might 
help the applicant. Visitors were also employed to call at the homes of the 
applicants to make careful investigations. The frequency with which the 
same names appeared in the lists of relatives of applicants and the family 
likenesses of individual applicants soon led to the conclusion that these 
paupers were one large closely related group and so received the name, 
“the Ishmaelites,” from the Ishmael family, the name of the central and 
worst family of these paupers. It was at this time that the Rev. O. C. 
McCulloch became interested in these people as mentioned above, saw the 
similarity of them to the Jukes, noting, however, the difference that this 
large set of people came from several hundred different family heads whereas 
the Jukes had all come from the one mother with five daughters. 

In 1888, McCulloch read a paper at the National Conference of Charities 
and Corrections at Buffalo, presenting a diagram where thirty different 
family groups of paupers had been traced out covering about two hundred 
and fifty separate families or households. He says, 


400 


ARTHUR H. ESTABROOK 


The central family—that which gives its name to the Tribe of Ishmael—first appears 
in Indianapolis about 1840. The original family stem, of which we have scant records 
as far back as 1790, is then in Kentucky, having come from Maryland through Penn¬ 
sylvania. John Ishmael married a half-breed woman and came to Marion County, 
Indiana, about 1840. He was diseased and could go no further. Three of his sons 
married three sisters from a pauper family named Smith. 

Since 1840, this family has had a pauper record. They have been in the almshouse, 
the House of Refuge, the Woman’s Reformatory, the penitentiaries and have received 
continuous aid from the township. They are intermarried with the other members of 
this group and with two hundred and fifty other families. In this family history are 
murders, a large number of illegitimacies and of prostitutes. They are generally diseased. 
The children die young. They live by petty stealing, begging, ash-gathering. In summer 
they “gypsy” or travel in wagons east or west. In the fall they return. They have been 
known to live in hollow trees or the river bottoms or in empty houses. 

In this sketch three things will be evident: First, the wandering blood from the half- 
breed mother; in the second generation the poison and passion that probably came with 
her. Second, the licentiousness which characterizes all the men and women, and the 
diseased and physically weakened condition. From this result mental weakness, general 
incapacity and unfitness for hard work. And, third, this condition is met by the benevo¬ 
lent public with almost unlimited public and private aid, thus encouraging them in this 
idle, wandering life, and in the propagation of similarly disposed children. 

Thus McCulloch tells of the conditions in 1888. Soon after this report 
McCulloch died and the study stopped at that point. Some of the original 
data have been lost; the official records are mainly extant. About 1915, 
the Eugenics Record Office took up the study of the Tribe of Ishmael at the 
point where it had been dropped at McCulloch’s death and the work 
has been carried on to the present except during the time of the participa¬ 
tion of the United States in the World War. 

The present investigation has shown that the different families of the 
Tribe came to Indiana, separately in most cases, on the general tide of 
immigration west from the original thirteen colonies along the seaboard just 
following the War of the Revolution. The early immigration into Indiana 
was mainly from southwestern Ohio and Kentucky. These people in turn 
had come either from the Carolinas through the Cumberland Gap or 
Tennessee or from Virginia, Maryland and Pennsylvania, overland by way 
of the Potomac River, over the mountains and then down the Ohio River. 
The make-up of the population of Virginia in Colonial days gives a clue to 
the origin of the Ishmaelites of Indiana. Labor was scarce in Virginia at 
that time and the Virginia Company to fulfil its contract brought to this 
country “idlers” and members of the classes in England who were “charge¬ 
able, dangerous and troublesome to the State,” and youthful vagabonds 
and later convicted criminals, some political but more the common, the 
anti-social, these latter being sent here to serve out their terms and then to 


THE TRIBE OF ISHMAEL 


401 


be set free in this country. After 1650, the deportation of confirmed crimi¬ 
nals, i.e., felons, to the American Colonies was a common practice. 

That not all of these anti-social persons sent to the Colonies were men is 
shown by an entry, one of many, of a ship leaving Lieth, England, in 1692 
“for Virginia, holding 50 lewd women out of the houses of correction and 
30 others, who walked the streets after ten at night.” This deportation 
continued until about 1770 but there are no figures as to the number who 
came here. In the lists of criminals that are now extant, some of the fam¬ 
ily names of the Tribe are found and very many more names are found in 
the lists of servants, i.e., people indentured or sold, because of criminal 
acts in England, for a term of years to planters. Although no actual lineage 
connection between these individual names and the Tribe family heads has 
been made the fact is interesting and suggestive of much. It is reasonable 
to assume that some of the Tribe families have gone back to these paupers, 
criminals and prostitutes sent to the Colonies. Some of the Ishmael names 
are very uncommon, which fact aids in this conclusion. 

It is estimated that the Tribe numbered six thousand people in 1885, 
coming from about four hundred different family heads. Today no estimate 
of the number of Ishmaels can be made because many of the lines of de¬ 
scent have been lost and so cannot be traced, but the number would not be 
less than ten thousand. They are now found mainly in Indiana, Kentucky, 
Illinois, Ohio, Michigan, Iowa and Kansas. 

There are three outstanding characteristics of the members of the Tribe; 
pauperism, licentiousness and gypsying. In a paper such as this, statistics 
and family histories are of little value to tell the story and are out of place. 
Cases could be cited endlessly where a family for generation after genera¬ 
tion have been professional beggars and paupers, receiving both public and 
private relief. The names of these families are found year after year on the 
township trustee books and soon the children, now as parents, appear under 
their own names, asking for help. In the same way the professional beggars 
tour the town both in the residential and business districts with always a 
pitiful tale, which never bears investigation, sometimes the eyes washed 
with bluestone water to make them inflamed and the individual claiming 
blindness, or an injury to the hand or foot, kept irritated for months as a 
plea for help and an excuse for unemployment. There is hardly a family 
in the Tribe that has not had some beggars; in many of the families almost 
all the members are adept in the art of begging and have plied their trade 
for years. They always carried a basket, sometimes with dirty soap or 
bluing as a pretext to call, but the basket was ready to hold anything that 
would be given, either clothing, food or money. As blindness has been 


402 


ARTHUR H. ESTABROOK 


quite common in several families of the Tribe, a familiar sight had been a 
blind man or woman led by others begging from house to house. Young 
children too have been taught to beg. The amount of the actual relief or 
help secured by the Tribe far exceeds anything given to the Jukes or the 
Nams. No estimate of this can ever be made. When the struggle has been 
too hard often indoor care for members of the Tribe has been furnished in 
poor asylums or other institutions. Many have spent the last years of their 
lives in almshouses. It has not been uncommon for three generations in 
one family to be in the poor house at the same time. 

Licentiousness is perhaps the next interesting trait which has charac¬ 
terized the Tribe. The loose marriage relationship has been one of the out¬ 
standing features. A fair proportion of the marriages in the Tribe have 
been according to the law, both parties legally free, a license secured and the 
marriage ceremony performed either by a clergyman or a justice of the peace, 
with the two persons concerned remaining together until the death of one. 
There have been many cases where two people have merely cohabited under 
the common law marriage and children born under these conditions have 
been considered legitimate. Divorce has been very easy in Indiana. Many 
of the Ishmaels had no comprehension of or respect for the marriage vows 
and so their consorts were left or changed at will. One example of this will 
indicate the extent to which it existed in one family even though this is an 
extreme case. Four members of one fraternity, one man and three women, 
each had respectively, six, five, five, and seven marriages or matings; in 
no case death causing a separation, no divorce actions completed and only 
about half of the matings accompanied by a marriage ceremony. A daugh¬ 
ter of the woman who had seven marriages was herself a prostitute, married 
ten times, several of these by ceremony—no divorce between any two 
marriages. 

Prostitution as will be seen from the above was common in the Tribe. 
At one time the greater proportion of the women keeping houses of prostitu¬ 
tion in Indianapolis belonged to the Tribe. Several of these houses were 
famous in this region of the state. The elegance of a few of these houses of 
prostitution in the late sixties, seventies and eighties of this past century in 
Indianapolis is often recalled by older police officials and newspaper re¬ 
porters who were familiar with the inside of these establishments. The Ish- 
maelites often used members of their own families as inmates of these houses. 
In one a woman and her two granddaughters, while in several cases a woman 
and her daughters, comprised the personnel of the bagnio. Many other 
Ishmaelites frequented houses of prostitution merely as casual inmates. 
There were many homes among the Ishmaels where illicit relationships 


THE TRIBE OF ISHMAEL 


403 


occurred. Some incest has been found in the Tribe. No figures as to the 
number of people married, divorced, children born in or out of wedlock, etc., 
are presented because of the lack of official registration covering these 
points. It can be said that the illegitimacy is greater than in either the 
Jukes or Nams. 

The other marked characteristic of the Tribe has been the wandering or 
“ gypsying” as it is called by the Ishmaelites. The earliest known of these 
“ American gypsies” as they were called, was John Ishmael, who had come 
to Indianapolis from Kentucky about 1825. In the next few years he made 
several gypsy trips towards the Ohio River and Cincinnati. As this country 
became settled these trips turned to the north part of Indiana into the Indian 
reservations. These reservations covered over fifteen hundred square 
miles and were open only to Indians but these gypsies, much more numer¬ 
ous by this time, seem to have been permitted at all times to enter them 
to hunt and fish. Here these people gypsied during the summer months, 
returning to Indianapolis for the winter. It was upon these trips that many 
of the early marriages of the Tribe have taken place. After 1845, northern 
Indiana was being settled rapidly and the Indians were gradually being 
pushed from the reservations. The gypsyings therefore ceased in this di¬ 
rection and started out again in a land less highly developed. This time 
the route was to the plains of Illinois and sometimes as far as Iowa and 
Kansas. 

Ordinarily these gypsy ings began in the spring and ended when “ roasting 
ears were ripe.” Sometimes however a family did not return in the fall but 
remained away that winter, returning to Indianapolis the next fall. When a 
family remained away over winter it is probable that it was most often 
spent in some county poor asylum. The asylum records in various coun¬ 
ties in Illinois and one record in Iowa show that some of these winter 
sojourns were in those poor houses and thus the worst of the winter was 
tided over. 

In the early spring the Ishmaels and many of the other related families 
put all their worldly possessions in a hand cart and started across the coun¬ 
try; later they were able to acquire an old broken down horse or mule, often 
a retired street car horse too old to use in that service, and a more dilapidated 
wagon in which to go on their wanderings. They would go often with 
several “wagons” in a party, sometimes alone. They camped in creek 
bottoms, near a settlement if possible, and lived off the country, begging and 
stealing. When they became tired of the place or, as often, were told to 
move on by the people who could no longer stand their depredations, they 
travelled on to the next place to do the same again. The residents along 


404 


ARTHUR H. ESTABROOK 


these routes still remember the gypsies; though they can call few by name. 
There were some places on these routes where the gypsies particularly liked 
to stay. At the first approach of cold weather the wagons were turned to¬ 
ward Indianapolis and they hurried back so as to get located in the city be¬ 
fore winter set in and as it has been expressed “ to get their names on the 
trustee’s books before frost appeared.” 

As whole families went together, old and young, parents and children, 
sometimes three generations at once, the tricks and habits of the road were 
passed on from one to the other. Several wagons would often travel to¬ 
gether and gypsies would then meet other travellers and in this way [mat¬ 
ings between the gypsies took place. Propinquity and like mating to like 
were the two factors controlling the matings in these groups. 

Another group of the Ishmaels were the so-called winter gypsies. This 
group, much smaller in numbers than the other, went south from Indian¬ 
apolis into southern Indiana at the beginning of fall, in wagons, swapping 
horses, gambling and living as best they might. These people had no par¬ 
ticular routes of travel; they simply went where they listed, returning to 
Indianapolis in the early spring. Many of the laborers in these families 
worked in the brick yards and so were unemployed during the winter. 
This winter gypsying did not continue long and does not seem to have been 
carried on regularly by those who did “winter gypsying.” These families 
were of a slightly higher mental level than the summer gypsies. 

Such is the picture roughly and briefly of the Tribe of Ishmael. The 
individuals in this large group of feeble-minded folk are continuing to mate 
like to like, and are reproducing their own kind. Some few branches of the 
Tribe have mated into better stocks, but these are so few that they are hardly 
noticeable. The few placed in orphans’ homes and new environments 
have in some cases done better, but this has not changed the whole mass to 
any extent. The greater portion are still the cacogenic folk as found by 
McCulloch and are breeding true to the type. These germ plasms have now 
spread through the whole middle west and are continuing to spread the anti¬ 
social traits of their germ plasm with no check by society. The story of 
the Tribe of Ishmael is but another picture of the Kallikaks, the Nams and 
the Jukes. 


HERITABLE FACTORS IN HUMAN FITNESS AND THEIR 
SOCIAL CONTROL 

WILHELMINE E. KEY 
Race Betterment Foundation, Battle Creek, Michigan 

The foundations of national power are, in the last analysis, biological. 
We no longer look upon the life of a people as something unfolding itself in 
a manner wholly mysterious, and then as certainly going down to decadence 
and death. The growing clearness with which the progress of science en¬ 
ables us to view all life phenomena has thrown light upon human develop¬ 
ment in its physical, mental and moral aspects. Increasingly, we are being 
won to the view that the elements of a nation’s strength he in the inherent 
traits and tendencies of its people. Passing events, as they crystallize into 
history, are but the interplay between such traits and tendencies and various 
external influences, chiefly those expressed through the science and art of 
the period. Breeding from the best elements of the population brings a 
growing preponderance of those best fitted to take highest advantage of 
these influences. But science and art, in so far as they are not borrowed, 
are of themselves the abiding effect of this breeding from the best. Since 
their refinement of method too often encourages the survival of the unfit, 
the question of a nation’s continuing integrity resolves itself into a nice 
balance between certain genetic and social forces which inhere in the blood 
lines of its constituent peoples. 

The chief concern of the eugenicist is then the production of better human 
strains. This involves the physical problems of heredity as conceived by 
the geneticist. But vastly more it involves the social and moral reactions 
as conditioned by our physical and mental constitution in relation to the 
complicated organism known as the social order of today. We thus glimpse 
at the outset the complexity of our problem and the necessarily one-sided 
treatment which any brief study must give. 

We none of us need to be told how complex is the thing we call human 
personality. Still most of us are aware of outstanding capacities and domi¬ 
nating tendencies in conduct. In the analysis which follows, certain capaci¬ 
ties and tendencies of this kind were selected for intensive study and their 
occurrence noted from generation to generation in relation to types of 

405 


406 


WILHELMINE E. KEY 


marriage which the individual was able to make. These capacities, notably, 
calculating ability, planfulness and perseverance were evaluated by means 
of the social test, that is the test of efficiency in the activities undertaken. 
Modern psychology has developed tests which, no doubt, are far superior 
to the social test. But in regard to these laboratory tests two things are 
to be observed. Their findings so far have coincided with the results of 
observation and experience on success in life, that is efficiency in activities 
undertaken. We are therefore justified more than ever in placing reliance 
on the social test, especially since it is obviously impracticable to apply the 
laboratory tests to all the members of the families we are studying. 

Intensive study of such family networks shows widely differing poten¬ 
tiality of their various lines. In the Rufer Family, described at length 
in Carnegie Publication No. 296, five widely differing lines were discovered. 
This family was founded in the latter part of the eighteenth century by an 
able, industrious, plucky pioneer and his easygoing, amiable wife, who, it 
appears, was totally lacking in sense of number and proportion. Present 
day representatives of this family network show widely divergent social 
status. They spell their names differently and resent the implication of 
relationship. It was the custom of Aaron Rufer to count out every morning 
the potatoes needed for the family dinner; this done, his wife knew enough 
to put them over the fire, when the sun was high, and cook them till they 
were done. She neither spun nor wove, and the rest of her housekeeping 
was on a par with her cooking. There are many Mary Rufers to be found 
in disordered homes in a certain county of western Pennsylvania today. 
They are all descendants of Aaron and Mary in lines where marriage has 
consistently been into strains showing similar lack. In three of the lines, 
by contrast, there is conspicuous presence of planfulness, perseverance and 
ability with number; and here marriage has almost uniformly been into 
strains showing these abilities. 

Lines C and D are particularly interesting. In line C there is complete 
absence in later generations of anything that might be called social inade¬ 
quacy. It has realized the half-playful suggestion of a eugenicist of another 
age, that with proper selection (the right sort of marriage) a defect may be 
blotted out of a family just as “a careful gardener blots out the blemish 
from a flower.” Line D, on the other hand, is replete with drunkards, 
prostitutes, petty thieves and paupers. When we go back far enough, we 
find this astonishing divergence arising as a very slight difference between 
two sons of Aaron and Mary. But the slight balance in favor of the better 
endowed son brought about marriage with a better woman who belonged to 
a strain superior to her husband’s as well as superior to that of her 


HERITABLE FACTORS IN HUMAN FITNESS 


407 


sister-in-law. Line E, founded by Aaron’s imbecile son through his marriage 
to a sexually lax girl, for whom well-meaning neighbors wanted to find a 
home, has been persistently imbecile. Never but once in five generations, 
so far as could be determined, has there been fertile union with a member of a 
good family. This resulted in an illegitimate daughter whose two children 
are apparently normal. All the rest are mid-grade imbeciles, fortunately 
in the custodial care of the state. This extended experiment in mating then 
teaches the value of matings with good stock in establishing socially fit 
lines. 

The study reveals too many of the social factors involved in the preserva¬ 
tion of socially valuable traits. The mere possession of the traits in question, 
that is planfulness, perseverance and ability with number, has meant union 
with strains having similar abilities (assortative matings) while their ab¬ 
sence has brought about union with individuals having the same or differ¬ 
ent defects, and this has operated to accentuate the condition of degeneracy 
and defect. Studies with the quantitative variation of these abilities point 
to a segregation of factors to produce on the one hand increase, on the other, 
progressive degeneration with reference to the traits in question according 
to the type of mating made. 

Such findings attain grave significance when we consider the great fecun¬ 
dity of degenerating lines, their proneness to draw to themselves the weaker 
individuals of other mixed strains to form centres of degeneracy and defect. 
Every eugenic worker is familiar with such centres. The certainty with 
which they flourish in defiance of all attempts at amelioration, often because 
of such amelioration, emphasizes as one of the chief functions of our institu¬ 
tions for the care of the defective and diseased, a sufficiently detailed study 
of the family history of their patients to determine the blood lines from 
which they come. Only in this way can we give genetic potentiality its 
due weight in deciding the fate of such patients. Under present conditions 
of economic stress, the superintendents of these institutions are being urged 
to make the persons under their charge industrially efficient and insufficient 
appropriations'for maintenance are adding their share to tip the balance 
toward the wrong side, that is in favor of early discharge and the almost 
certain marriage and multiplication of the unfit. When considering ap¬ 
propriations for this type of study we should always remember that as a 
people we cannot afford to do anything but spend all that is needed to cut 
off degenerating lines. 

Again, public sentiment in many parts of our country is still so lax or 
uninformed that individuals belonging to many such tainted blood lines 
have not yet been committed to any institutions. With one exception, the 


408 


WILHELMINE E. KEY 


dozens of degenerates before described as belonging to line D of the Rufer 
family are all at large. Following them is like following the trail of the 
serpent. Inevitably they lead to the dregs of the community, or to knots of 
human derelicts, a moral menace and economic drag. The work within the 
institution should be supplemented by surveys, federally or state supported, 
and appropriate preventive measures taken. 

Taking up a different phase of our question, we come to the abilities 
known as the aptitudes. These have also been shown to follow blood lines. 
If we study their occurrence in moderately gifted individuals, your family 
and mine, I am sure we can all easily be convinced of this. Moreover, 
these blood lines are found to move up or down in accordance with the type 
of marriage made. Peculiarities of reaction of what appears to be in the 
main a similar makeup placed in a varying or widely differing environment 
furnish interesting examples of the so-called greater effect of environment 
in determining social effectiveness. There is here a wide field. We must 
cease to debate the futile question of heredity versus the environment, and 
set ourselves to evaluating abilities, so far as we can, and analyzing the 
interplay of extraneous conditions on these innate capacities. The in¬ 
vestigation of the social and economic misfits is peculiarly valuable. In 
this work, the physician, the teacher, the enlightened parent are showing a 
promising tendency to co-operate with the specialist in casting up the ele¬ 
ments of a given situation. It would seem that here again there lurk 
certain dangers. One is the danger already touched on. That is that all 
these refinements will eventuate in the survival of the unadaptable or at 
least those difficult of adaptation. Our development should tend toward 
increasing power of self-adaptation. And, accordingly, should not these 
refinements of means be paralleled by a checking of reproduction when the 
difficulties of adaptation threaten to become too great? Checks in such 
cases might well be largely voluntary, self-imposed through eugenic educa¬ 
tion, as when a highly unstable individual, who is still able to carry his own 
weight, remains single in preference to marrying when marriage into a 
strain threatened to accentuate that instability. 

Passing to the observation of assortative mating in yet other fields, we 
come to the production of those personalities which have given us our 
peculiar development in industry, science and art. These personalities 
have, through the study of their biographies and genealogies, been found to 
take their rise in the following manner. They have apparently arisen by 
the fortunate crossing of able lines in conjunction with a social or economic 
opportunity which favored the development and the play of their peculiar 
gifts. Sometimes, they even establish the favoring conditions instead of 


HERITABLE FACTORS IN HUMAN FITNESS 


409 


waiting for opportunity to knock the proverbial single time. Occurring 
with such eminent individuals in the blood network are usually found many 
others having similar abilities, or at least occupations which suggest similar 
trait-complexes. The majority of these may indeed be more or less obscure, 
still we find here and there what might be termed concentrations of ability 
of various types. We find one line remaining tillers of the soil, others are 
given to merchandizing, while still others may show marked artistic trends. 
In the last-named cases, the artistic ability may “shade off” into a well- 
marked artizan group. Again we find a more or less well-marked altruistic 
bent for many generations, the latter as J. McKeen Cattail has shown evinc¬ 
ing itself in one generation in the selection of the ministry, in the following, 
when preaching has lost some of its power and prestige, in preference for the 
career of teacher. 

In the same way we find pioneering to be characteristic of many families, 
inventiveness in others, such as the Fairbanks, Herreshoff and Lake families, 
and executive ability a distinctive trait-complex in still others. Our his¬ 
tory furnishes, too, many illustrations of signal ability in finance extending 
over three or four generations. In this connection, the names of Astor, 
Vanderbilt, Morgan, and Rockefeller will occur to all of you. It is true 
that the peculiar trait-combinations which make for this ability are differ¬ 
ent in the families we have mentioned, but in every case, the salient traits of 
the leading personalities find abundant representation in the network to 
which he belongs. Thus the story of our Americanism becomes the story 
of concentration of genetic potentialities of various kinds, fostered by 
favoring physical, social and economic conditions. 

Extending our view to the field of statecraft, education and reform, we 
again find it to be the story of germ-plasms chosen through the exigencies 
of the place and time. Types of leadership as exemplified by the Beecher, 
Abbott, Lowell, Edwards and Adams families, to cite only a few of the 
many possible instances, have arisen in the same manner as that already 
outlined. Detailed analyses of such eminent men as Jefferson, John Adams, 
Benjamin Franklin have been made by the writer and the traits which con¬ 
tributed to the abilities in every case found to have abundant representation 
in other members of their families. Usually the peculiar trait-combination 
which each possessed has been found to occur but once. We have, however, 
in Theodore Roosevelt an instance where they appear to have occurred 
three times. Archibald Bulloch, his maternal great-grandfather, and Robert 
Barnwell Roosevelt, his father’s brother are frequently cited as prototypes 
of this most illustrious “first American of his time.” 

In common with many great men, Roosevelt was inclined to minimize the 


410 


WILHELMINE E. KEY 


influence of heredity. He was proud of the fighting stock from which he 
sprang but beyond this gave little weight to ancestral factors. Time 
forbids the fascinating task of analyzing Roosevelt’s character and family, 
the result of months of intensive study. Suffice it only to say that when we 
survey the concentration of gifts bestowed from both sides of Roosevelt’s 
family, it seems almost inevitable that his course should have been what it 
was. Added to this we have the peculiar conditions, industrial, social and 
political playing their part in bringing out Roosevelt’s salient traits. And 
furthermore, the characteristic moral and intellectual reactions of his family, 
were in him associated with a physiologic condition, present also in his 
father and others of his family which made it possible and even necessary 
to follow out every undertaking with the utmost intensity. What more 
natural than that in the political and industrial conditions presented by our 
country as a result of its phenomenal growth such a forthright, courageous 
nature as Roosevelt’s, loving power and conflict, but loving the common 
people more, should find here a fertile field for the highest expression of its 
powers? It is this action and reaction between his restless, irresistible 
energy, playing with a wide range of intellectual tastes and powers and his 
sympathy with the fundamental tendencies of the American people that 
make of him such an outstanding example of the creative influence of a 
personality in our national life. 

Hitherto the social and genetic process just outlined has appeared to 
take care of itself. There are, however, grave indications that a change is 
setting in. The dying out of many highly gifted lines and the low fecundity 
of others, which would, if producing larger families, increase their chance of 
producing gifted individuals are among the signs of this change. The 
salient economic and social factors involved in this grave situation have, 
most of them, still to be evaluated. We have already touched on several 
of them in the course of this talk. Among other means of correcting the 
anti-eugenic influences now at work to undermine our integrity as a people, 
we might name the following: 

1. A return to saner and simpler standards of living such as will 
make possible the rearing of larger families in our “middle class.” This 
would eventuate among other things, in a “back to the land” movement 
with all that that could entail for the well-being of the generations to come. 

2. A campaign of education among all classes of our population which will 
foster the eugenic conscience. Many of our people, notably our young 
women, products of our so-called higher education, have been victims of 
the “ingrowing eugenic conscience.” While undoubtedly the race has 
profited by the dying out of these rankest expressions of our rampart 


HERITABLE FACTORS IN HUMAN FITNESS 


411 


individualism, in other cases, their failure to become parents has meant a 
distinct national loss. 

3. The encouragement on the part of our numerous foundations, not of 
celibacy and comparative sterility in their members, but of marrying and 
rearing families having the desirable genetic potentialities. While this 
might result in some loss in the present generation, it would surely be com¬ 
pensated for in time to come. This brings us to another important factor. 

4. The concept of service in contrast to that of personal aggrandizement. 
Our increasing socialization is bringing as a natural sequence such a con¬ 
ception of responsibility to the state and to humanity in the type of marriage 
made. This is the responsibility of maintaining through selective matings 
of the best available the highest possible level of ability in the family. 

5. Early and more adequate economic adjustment to render possible early 
marriage and parenthood. Our hospitals and asylums are filled with the 
victims of maladjustment. A fair proportion of our delinquents are ex¬ 
amples of maladjustment which might and should have been prevented. 
The writer has in mind a youth of fifteen who was admitted to a Training 
School for Feebleminded on a charge of incorrigibility. He had been a 
serious problem to his family since he came to them from Italy two years 
before. Had failed to get on at school, ran the streets, frequently climbing 
telephone poles and ranging about like wild, so that he was followed by a 
hooting crowd of boys who thought him crazy. Apprenticed to a tailor, he 
had continued his errant ways till the authorities could find no means of 
control but a school for the feebleminded. Here, fortunately, he found 
those who understood him, and, under their influence, he became gentle¬ 
manly, eager to learn and a general favorite among his teachers and mates. 
He showed marked ability in wood working, sang well and played the violin. 
Mental examination found him to possess adult mentality (his teachers had 
put him in a class of five-year-olds) and study of his family history revealed 
the springs of his abnormal conduct to be simply these: He came from a 
family which for generations had been artisans and small farmers; until he 
came to America, he had freely roamed his silent native hills. Is it any 
wonder that shut up in the confines of a tenement, in a narrow turbulent 
street, put into a school where all his excellences were ignored, his faults 
exaggerated, his efforts ridiculed he should have acted as he did? Removed 
to a sympathetic environment he made wonderful progress and is now sup¬ 
porting himself and his family at his chosen line of work, the making of 
furniture. Is it not a shameful commentary on our school system that this 
boy should have had to find his way into a training school for feebleminded 
before his peculiar abilities were discovered and fostered? 


412 


WILHELMINE E. KEY 


The least we can do is to apply mental tests to determine the mental level 
of all children in our schools. But at best, the mental test furnishes only 
what may be called a cross-section of the pupil’s abilities at a given time. 
The results of these tests should be supplemented by family history studies 
to determine the trend of mental and emotional development. This is 
especially valuable in cases where the child or youth shows delinquent 
tendencies or bids fair to become an unusually brilliant student. In such 
cases, the ounce of proverbial prevention is trebly worth the pound of cure. 
How many of our young people, now irretrievably lost, might not so have 
been saved to become worthy and even able citizens? We want the survival 
of the self-adjusting; but we may well pause here to inquire whether our 
present social order is the one ideally fitted to encourage the adaptation and 
survival of the best. We can get along, in the writer’s opinion, with less 
push, particularly when that is combined with unscrupulousness, and should 
work toward a condition where other and more ideal excellences abound. 
Our eugenic philosophy is as yet in its mere beginnings. These are only a 
few of the measures, which as good Americans we must compass if we would 
preserve lines of socially fit families and perform the desired service to 
national and racial growth. 


A STUDY OF ONE HUNDRED AND FIFTY ADOLESCENT 
RUNAWAYS 

ELIZABETH GREENE 
Waverly House , New York City 

The hereditary factors in crime have long been an interesting problem for 
eugenists. If such factors do exist, possibly the best place to study them 
will be in the adolescent delinquent, who is just beginning to show anti¬ 
social tendencies. It is these young offenders with whom the New York 
Probation and Protective Association does its work, girls who are just 
beginning to harass their parents and guardians with poor work habits and 
late hours, girls who have just entered into their first illicit sex experience, 
who have taken something from their employer, or more especially girls 
who have revolted against the discipline of home and have absented them¬ 
selves without the consent of their parents, that is the so-called runaway 
girl. 

It is this group which I want to bring to your attention, because it is the 
largest group which we handle, because it is a group with clearly marked 
inherited traits and one which is played upon by eugenic and disgenic 
forces. In thirteen months (May 1, 1920, to May 31, 1921) 150 runaway 
girls passed through our Mental Clinic. Some were found in the homes of 
friends old or recently acquired, some in railway stations, some in furnished 
rooms. Policemen, detectives, social workers brought them in. We studied 
them, planned for them and sent them home. Of the entire number, 51, 
or about one-third came from homes in the city. As many of these were 
persistent runaways and had remained undiscovered for long periods, we 
must consider them runaways, although the distance travelled was incon¬ 
siderable. The remaining 99 came from fifteen different states, more fre¬ 
quently, of course, from neighboring ones. Still, 7 came from Delaware, 
Maryland and the District, 8 from Ohio, 3 from Michigan, 1 from each 
Kentucky and Missouri. 

The girls were all young, largely because running away is closely con¬ 
nected with the restlessness and dissatisfaction of youth. It is the curiosity 
and self-confidence of the girl of sixteen which brings her to the city, 
confident that she can take care of herself. The youngest whom we received 

413 


414 


ELIZABETH GREENE 


was only fourteen, while the largest group of all were the sixteen-year-olds. 
If we use the statistical device of the frequency table, we find that the first 
quartile falls in the seventeen-year-old group, and both the second and third 
quartiles in the eighteen-year-old group, showing that as far as age is con¬ 
cerned the group is fairly homogeneous. 

When we consider the work histories of these girls we find that all but six 
had been employed, the exceptions being due to extreme youth. In fact, 
one little girl came to us from New Jersey with her school books under her 
arm. Sixty-one (or 41 per cent) of the girls were factory workers, thirty 
(or 20 per cent) clerical workers, twenty-four (or 16 per cent) domestics, 
sixteen (or 11 per cent) mercantile workers, while the remainder fell in 
small groups, telephone girls, dancing teachers, and show girls. They 
differ from the prostitute group in that there are fewer domestics and wait¬ 
resses among them. There is probably nothing else particularly significant 
about this distribution. As adolescents they tend to find employment 
only in unskilled and uninteresting work; they become bored and run 
away. 

The mental diagnoses were arrived at by means of a complete physical and 
psychiatric examination, as well as a mental test. Seventy-eight, or more 
than half of the girls, fall in the normal or full normal groups, about a sixth 
are borderline, and another sixth defective, the remainder being constitu¬ 
tional inferiors, psychotic and psychoneurotic. We find that runaways 
grade higher mentally than girls brought in for sex offenses. 

The nativity of these girls is of especial interest. Twenty-two per cent 
were born in foreign countries and came to this country with their families. 
When we consider the nativity of the parents, we find that 57 per cent of 
these girls are the children of foreign-born parents, 28 per cent the children 
of native born parents and 13 per cent the children of mixed parentage. 
When we compare these figures with those in the 1910 census for women in 
New York state we find that a larger proportion of the runaway girls are the 
children of foreign parents than of the general female population. That is, 
they are the direct descendants of persons who themselves left home to 
better their conditions or see new places. 

We have, therefore, 150 adolescents, all young, many the children of 
foreign born parents, trying to adjust themselves in work for which they 
have had no training, and to parents whose ideas are very different from 
theirs. The way out is to run away, although usually this is only attempted 
after frequent quarrels, and constant criticism. The act is in itself depend¬ 
ent not only on the social situation but also on the personality traits of the 
individual girls. 


A STUDY OF ADOLESCENT RUNAWAYS 


415 


A few of them are no doubt typical wanderers, such as Dr. Charles Daven¬ 
port has described. A little girl who suggested this type was an Italian 
who was brought to us after her third absence from home. The parents 
were immigrants, while the maternal grandfather had left Italy for South 
America as a young man. A little better example was a girl who had been 
placed by the Juvenile Court as maid in a Connecticut city. She had very 
little amusement and one day when her mistress left her to get lunch and 
wash a tub-full of clothes she ran away to New York. Her family history 
would delight any eugenic field worker. Both parents and a maternal aunt 
were alcoholic, while an older sister had been committed to a school for girls 
in Connecticut. Moreover, the family proved to be well-known, both in 
Utica and Hartford, as wanderers; both parents were peddlers, while the 
uncles were horse traders and troubadours, their sisters and wives wandering 
with them. 

Besides wanderlust we find in these runaway individuals the native push 
and energy which mark the hyperkinetic. Others show an entire lack of 
these qualities and are the so-called inadequates. That these two types 
may do the same thing from quite different motives was shown in three 
runaways who came in May. These little girls, the oldest seventeen, 
were idling in the square of a New England town when an interurban furni¬ 
ture truck stopped and the drivers offered them a ride. One, a round- 
faced, Anglo-Saxon, who had already lived in three New England states, 
accepted readily. She was delighted with the chance to see New York. 
The second girl, a moron, whose father is an epileptic and whose family is 
“on the town,” accepted because she had not the intelligence to decline. 
The third left home because her alcoholic father was so “distant and dark” 
when sober. 

In concluding we might say that one of the social problems of the adoles¬ 
cent is her tendency to run away. Runaways are always present in large 
cities, in stations, dance halls, and boarding homes. They show a distinctly 
American tendency to manage their own affairs at an early age and an adoles¬ 
cent dissatisfaction with their homes and revolt against parental authority. 
As the children of immigrants they are tied to no community by the associa¬ 
tions of generations, through faulty Americanization they have lost touch 
with their own parents. Most fundamental of all, however, are the in¬ 
herited traits which furnish the push and energy for journeys and the Wan¬ 
derlust which makes them second nature. 


MATE SELECTION 


ROSWELL H. JOHNSON 
University of Pittsburgh, Pittsburgh , Pennsylvania 


Mate selection or sexual selection as it was designated by Darwin is best 
defined as the production of racial change resulting from an hereditary 
difference between the mated and the unmated, or a correlation between 
mates. The subject does not lend itself to classification wholly by direct 
division and subdivision, but is better classified independently on the basis 
of various criteria. One instance may then come under several heads of 
such a classification. The following plan is proposed. 


Optimal 


Classified as to correlation 


I Periodic 
[Secular 


Assortative 


In assortative mating the phenomenon of interest is that likes tend to 
mate with likes. This has the effect, even where all the individuals mate, of 
accentuating attributes and so increasing the variation of the species as a 
whole, as compared with what would be the result if mating were wholly 
at random. The existence of assortative mating in man is quite evident, 
most particularly between those of musical interests, although needed com¬ 
parative statistical studies are as yet lacking. Its effect is fortunate for it 
has accentuated high attainments in certain lines, and in cases where the 
characteristics are undesirable, they have been concentrated in fewer cases 
more readily located and dealt with. 

In optimal mating, selection of mates is more frequent from those possess¬ 
ing a certain degree, or the nearest approach thereto, of some attribute with 
the result that the species in the course of many generations is brought to 
that degree of that attribute. If this favored degree is at the mode of the 
species then there is no evolutional effect, the effect being merely to hold the 
species at this optimum. This has been called periodic by Pearson. If 
the optimum lies on one side of the mode, then the species is led in that direc¬ 
tion and the selection is secular, in Pearson’s language. The race may thus 
be carried to an optimum within the range of variation then existing, to one 

416 



MATE SELECTION 


417 


outside the range or indefinitely, when the optimum is either the minimum 
or maximum possible and lies outside the range. 


Preferential 

Classified as to means ■( 

Restrictive 


f Conscious 
[Unconscious 

[ Direct 
<j Responsive 
[Arranged 


[ By differential death rate of the sexes 
| By geographical and social segregation 
• By willful celibacy 
By sterility 
By retarded marriage 


In preferential selection there is a higher mating rate at the optimal 
degree, because there is a direct preference for such individuals as mates. 


SEXUAL 

CLASSIFIED as to CORRELATION 


SELECTION 
(OPTIMAL 


{ PERIODIC 


CLASSIFIED AS to MEANS 


[assortat/ve 

f,PREFERENTIAL 

[restrictive 

EXTRA-UM/TAL SECULAR 


f 


PERIODIC 



SIZE OF ATTRIBUTE 


,SECULAR 

r conscious 

ft UNCONSCIOUS 
DIRECT 
RESPONSIVE 
ARRANGED 
( BY COMBAT 

J BY SEGREGATION 
1 BY WILLFUL CELIBACY 
[by STERILITY 

_ EXTRA-LIMITAL SECULAR 

1NTRA -LIMITAL SECULAR T O A DEFINITE GOAL ~\\ T O ANJNDEFINITE GOAL 





In restrictive selection there is no direct preference but a corresponding 
result is brought about by the fact that by some means the range of choice 
is so restricted or controlled that irrespective of preference, sexual selection 
takes place. One means is by differential death rate of the sexes. A well 
known example of this means is in the head hunters of Borneo where fighting 
between the males reduces their number selectively on the basis of fighting 
efficiency. Properly this belongs under lethal selection, but as a borderland 
category, it is included here. Another example would be the segregation 
of individuals of differing sorts in different places because of the location of 

































418 


ROSWELL H. JOHNSON 


industries, racial distribution and the like, coupled with a varying ratio of 
sex to sex in different places. The conditions which produce a shortage of 
one sex will ordinarily do so in a selective way, so that the make-up of the 
population of that sex shows a different average of the attributes in question 
than if it equalled in numbers the opposite sex. 

Preferential selection may be conscious or unconsicous. By conscious 
selection is meant direct conscious discrimination. By unconscious is 
meant those cases where preference results from a competition in the at¬ 
tractiveness of potential mates in which the selector simply feels more 
“drawn” by one individual than by another without any analysis of the 
elements. The two processes may be mixed and there is doubtless a twi¬ 
light zone. 

During the course of courtship there is a change. At first preference is 
more purely a matter of weighing of attributes, but as the emotions become 
more and more involved, the analysis of attributes is so dulled as to give 
such truth as does apply to the old statement that “Love is blind.” As a 
result the process as a whole is by no means devoid of conscious discrimina¬ 
tive preference. 

Mate selection is “responsive” when an individual of one sex is stimulated 
to greater attractiveness in the presence of one individual of the opposite 
sex than before others. That this stimulation is in some cases in part 
manifested by increase of pulse is responsible for the conventional associa¬ 
tion of the heart with love. It is here that the woman has a more active role 
in courtship than is usually supposed. By responding more effectively to 
a certain man, this man is in time drawn to her more than others who do not 
evoke either her unconscious stimulation or effort to be attractive. This 
then on the part of the selected may be unconscious or planned. 

Preferential selection is “direct” when the mate prefers one over another 
candidate. It is here distinguished as “arranged” when the preference, 
whether conscious or unconscious, is by the parents, guardian, or friends, or 
even a broker. These may arrange the match directly or arrange to bring 
into acquaintanceship under propitious conditions the individuals whom 
it is desired to mate. 

The elements of preference are not all of equal desirability racially nor of 
equal interest eugenically. Some eugenists have shown greater concern over 
physical characteristics. This is not shared by the writer because with all 
possible humanitarian mitigation, lethal selection is still sufficiently active 
to prevent serious physical retrogression. On the contrary, to eugenics 
alone can we look for salvation from a serious decline in the mental and 
moral attributes of the race. 


MATE SELECTION 


419 


The r61e of physical attractiveness in sexual selection is of paramount 
effectiveness. In pointing this out, Knight Dunlap has gone further and 
expressed satisfaction that beauty is the leading element in mate selection. 
Beauty, however, embraces two classes of elements toward which our atti¬ 
tude must differ. In so far as good color, clear skin and normal weight 
for the size in both sexes and adequate width of pelvis in women produces 
beauty it need not be begrudged its effectiveness. But in reality such ele¬ 
ments of beauty are quite common and contribute only a small part of the 
effective ensemble. The other elements, while of great effectiveness, es¬ 
pecially in selection of women, are mainly merely details of facial shape and 
that distribution of adipose tissue most in vogue at the time. The great 
effectiveness of these feminine elements so stimulate a man that he neglects 
to allow, on the average, a proper weight for elements which vastly transcend 
them in importance. Therefore sharp issue is here taken with Dunlap in 
his contention that beauty consists mainly of items of important racial 
value and therefore should properly be the leading element in mate 
selection. 

Those mental traits that are most stimulating are the most effective. 
Hence vivacity leads. Doubtless the selection for vivacity was useful, 
but it may easily have gone too far in some strains. It is by no means a 
measure of mental efficiency and is frequently associated in its higher de¬ 
grees with instability and hyperthyroidism. Indeed it seems probable that 
much of the instability, neurasthenia and mental disease of today is the result 
of the relative over-effectiveness of vivacity in mate selection. Luckily 
in the selection of males vivacity yields more to a consideration of the 
fitness of the man as a steady provider. 

On the other hand, the non-stimulating qualities—stability, persistence, 
endurance, poise and judgment are undervalued traits. The young man 
especially, as he, more than the young woman, is too susceptible to mere 
physical charm, should be on his guard against the under-valuation of these 
traits. For every slight advantage in beauty and vivacity he obtains in 
his mate he is, on the average, foregoing a much larger degree of important 
excellences in other lines which are less stimulating. 

It is commonly alleged that men prefer to marry women of intellectual 
capacity inferior to themselves in order that their feeling of leadership may 
not suffer. This in my opinion is incorrect in the long-run, because mental 
alertness, fortunately, unlike some other mental attributes mentioned, does 
have a stimulating effect and hence is effective without the necessity of 
conscious preference. 

Public record of the high scores in the systematic mental testing now 


420 


ROSWELL H. JOHNSON 


becoming common would probably have a decided value in making more 
evident mental differences now hard to discriminate and recognized within 
too narrow a circle of acquaintances. 

The accomplishment of the sought and the accomplishment and longevity 
of ancestors, uncles, aunts, brothers and sisters may also well influence one’s 
judgment in mate selection. 

It is the fashion to decry the influence of family and wealth upon mate 
selection and of course the individual herself or himself should be the main 
consideration. There is, however, some degree of positive correlation be¬ 
tween social and financial status and the directly desirable qualities, so that 
we may be less concerned with these less personal qualities than by the 
over-emphasis of mere physical attractiveness. 

Repute in men, and to a less degree in women, is of high selective value 
and has enough correlation with desirable attributes to be of real eugenic 
value. Since the standards of desirability and the relative rank in repute 
of various attributes are subject to change, it is of moment to the eugenist 
to inquire into the causes of these changes. In general there is a competition 
between the groups superior in each of several traits who wish their particular 
excellence to contribute as important an element of repute as possible. This 
conflict goes on between those who glory in their birth either in the nobility 
or in an “old family,” those who wish appreciation of their wealth, and a 
third group of “intellectuals” not included in the other two groups. This 
last is sometimes subdivided into artistic, literary, scientific and “public 
spirited” groups and even into subdivisions of these. Each group yearns to 
be relatively highly appreciated. The repute accorded to these elements 
varies with time and place and is ever in a state of flux. Parallel with this 
“struggle” between groups of superiors is another effective between the su¬ 
perior and the mediocre, who decry, either directly or more usually by im¬ 
putation, generally unfair, of some questionable quality such as snob, 
high-brow, big-head, “aristocrat,” blue stocking, swell, “high and mighty,” 
pedant, “holier than thou,” googoo, and the like. 

These phenomena are more easily examined in undergraduate life, 
especially that of the interesting effort of mediocrity to avoid recognizing 
excellence. We have the mediocre students calling those who lead in class 
“ greasy grinds.” At Dartmouth the epithet “ wet” probably from the wet 
towel around the head originally synonymous with “greasy grind” has been 
extended to deride any brilliant student. One reason for the derogation of 
mental excellence is to permit a relatively higher repute for athletic prowess 
and for “camaraderie,” but it is largely an attitude of the mediocre for their 
own protection. 


MATE SELECTION 


421 


Since the majority are relatively mediocre and are all seeking solacing 
thoughts, the idea of underestimating the excellence of the superior will 
always be of great seductiveness. This makes more difficult the upholding 
of the best standards of preference in mate selection. 

Yet at different times and places society has shown itself capable of dis¬ 
proportionately high productiveness in one field of activity or another. 
This has been partly the result of the varying distribution of man’s approval 
of the different attributes for a greater share of repute. Thus oratory and 
athletics in Greece, military prowess in Rome, and the arts in the Italian 
Renaissance, and later scholarship and research in Germany and wealth 
in the United States hold in succession places higher in man’s esteem than 
had been the case. The significance for mate selection lies in the relative 
rank of the various elements of repute of the time and place under considera¬ 
tion and here social progress and racial progress are nearly hand in hand, for 
an excellent distribution of the elements of repute favors each. Indeed 
the best distribution for one is nearly the best for the other. 

One divergence only arises. One commonly hears it said that some 
specially gifted individual might accomplish more for immediate social 
progress if childless or possibly even if celibate. This has led a few “ femin¬ 
ists” to protest against the dictum of the eugenists that all superior persons 
should adequately reproduce. At this point the eugenist appeals confidently 
and urgently for the ultimate benefit of the species. This is best attained, in 
the long run, by the adequate reproduction of even the specially gifted; 
for their superior germ plasm will thus yield in the future proportionately 
greater social and intellectual contributions. This is because the long suc¬ 
cession of persons produced from that chain of germ plasm will make a 
greater social contribution than that possible by the one life because of 
its celibacy with a resultant destruction of valuable germ plasm. 

A unique feature of preferential mate selection has arisen in recent years 
by the introduction to our species of the method of artificial impregnation 
now well known and not infrequently practiced in the case of horses. The 
author is reliably informed that in one of the largest American cities many 
women whose husbands are sterile now resort to a certain physician who 
inseminates them artificially. As is natural, these women stipulate the race 
of the father whose identity they rarely know. In some cases they are 
further concerned with his quality and attributes. It is obvious that this 
physician has a very heavy responsibility in thus determining the quality 
of many of these offspring. If he will, he can render a very valuable eugenic 
service by declining to inseminate inferior women and by obtaining the best 
fathers possible. 


422 


ROSWELL H. JOHNSON 


Writers on mate selection have usually assumed that rectification and 
intensification of standards of preference are the sole objectives of eugenists 
in this field. This error arises from too intent a concentration on the anal¬ 
ogy of lower animals. In man especially in recent years willful celibacy 
and childless matings have become relatively important elements. 

Celibacy has varied in its causes with the years. Formerly it was mainly 
religious and such celibacy still persists, although it is, I believe, on the 
wane. Such celibacy has cut off two groups from parenthood. One was 
inferior because it contained those actuated merely by a desire for the pro¬ 
tection and physical inactivity afforded religious celibates and their easy 
life. The other group was superior because for a long period, and still to 
some extent, it has been composed mainly of the brightest and most idealis¬ 
tic boys in a family who were led to train for the priesthood. 

Once the churches now having the priestly celibacy realize how the suc¬ 
cess of their own cult is reduced by breeding away from the qualities they 
desire, its abandonment will command an ever growing sentiment. 

But it is the newer celibacy that is more distressing—that evidenced by 
the low marriage rate of many superior women. The evidence concerning 
woman college graduates has been easy to obtain and these may be taken to 
represent the group. In their low marriage rate we apparently have the 
joint result of two unfortunate difficulties—first, the under-valuation of 
mental ability and the over-valuation of mere physical attractiveness 
by men as elsewhere decried. Nearly as important is the inefficient 
standard on the part of these women by which they respond only to poten¬ 
tial candidates whose qualifications are made so high as to render their 
number so greatly restricted that the chance of marriage is seriously reduced. 
Mathematically we may express it as follows. A woman who ranks thir¬ 
teen where one is the most superior and one hundred the most inferior, who 
feels no mating interest in any man less than rank six reduces her chances 
of marriage to less than one-half and therefore acts dysgenically. Theoreti¬ 
cally such a woman while young may well have an excessively high standard, 
but as her mating years pass, her standard should step by step slacken so 
that she will marry eventually, provided only that the average quality of 
her mate is not actually so inferior as to make the average of the couple 
below mediocrity. In other words, a superior woman today needs to be 
more concerned with marrying and less with her standard of preference; 
unusual as that sounds from a eugenist. 

Direct admonition is of course relatively unimportant in modifying cus¬ 
toms, so it behooves us to examine the conditions prevailing to see if any of 
these are being modified or are modifiable. Some of the causes of this low 
marriage rate are: 


MATE SELECTION 


423 


1. Isolation of sexes by separation in their schools, organizations such as 
Y. M. C. A. and Y. W. C. A. and even social clubs. 

2. Segregation in their work largely with women and married men. 

3. Limitation of proper avenues for widening acquaintance, on the one 
hand in small towns and on the other in the large city. The war Com¬ 
munity Service did a great deal to bridge this difficulty temporarily and 
some of its work has left permanently surviving institutions and customs. 

Passing to a consideration of restrictive marriage, several elements which 
reduce the number of effective mates require consideration. 

1. Incontinent celibacy. 

2. Inherent sterility. 

3. Sterility the result of disease. 

4. Induced sterility. 

5. Prevention of conception. 

6. Retarded marriage. 

The incontinent celibate is usually the parent of few or no surviving chil¬ 
dren. This may be due to infection, sterility of the temporary mate, greater 
efforts to prevent conception, or in case of conception, greater readiness to 
abort. Furthermore, offspring of such a union are often still bom, or die 
prematurely because of syphilitic infection, so that concern with this class 
from a eugenic standpoint is mainly to keep superior young persons from 
finding their way into it. With the improvement of the prophylaxis and 
treatment of the venereal diseases, births will become more common from 
this group of incontinent celibates because sterility will be decreased. Such 
a condition will demand more rigorous attention to a feasible program of 
restrictive eugenics. 

Unfortunately, many otherwise desirable individuals are inherently ster¬ 
ile, nevertheless it is probable that on the whole there are more inferior than 
superior individuals in this group. Sterility unknown to the other mate at 
the time of marriage should be a proper ground for divorce for those who 
desire it. Sterility the result of disease may be classified into that resulting 
from venereal diseases and that from other causes. The latter may be 
passed as relatively unimportant. 

This former group may be divided into those individuals infected as the 
result of their own moral transgression and those innocently infected by non- 
sexual contacts or by infected mates. 

Those infected as a result of moral transgression will average inferior, to 
judge from the nature of the inmates of genito-urinary wards of hospitals as 
compared with the other wards, as I am informed by Captain Paul Popenoe 


424 


ROSWELL H. JOHNSON 


from his experiences in army hospitals. Venereal infection is particularly 
important in holding down the spread of some inferior races that would 
otherwise be very fecund. Should the venereal diseases ever be effectively 
treated or prevented, it would be very important that means of birth con¬ 
trol be diligently spread among the married couples of such races. 

As to those innocently infected, one concludes that they average inferior, 
from what is known of assortative mating. 

This class of sterility then on the whole is eugenic in its selectional effect. 

Permanent sterility without impairment of sexual functions can be pro¬ 
duced in each sex by operation and by other means and has been so pro¬ 
duced voluntarily by a few to permit more sexual freedom. Some eugenists 
have been concerned lest this practice might become so common as to 
endanger the birth rate. 

But since the sterilized, as a whole, would average decidedly inferior, at 
least in some important attributes and since the general birth rate could well 
stand some diminution, this practice does not at present seem inimical to 
race progress. 

It has been argued that a wider knowledge of the prevention of concep¬ 
tion will too generally result in childless or inadequate families among the 
superiors. The passing of laws against the dissemination of such knowledge 
is not the way to meet this difficulty, for such knowledge, long since common 
property in Europe, has already reached, in general, the superior group in 
American cities. Such knowledge is of such great eugenic usefulness when 
once thoroughly widespread that the suppression policy should be aban¬ 
doned, even though for a time population might cease to increase. Our 
effort in this connection should be made to distribute its application on a 
eugenic basis. 

Since a retarded marriage often reduces the number of progeny and 
always lowers the. rate of increase per century, the effectiveness of such 
matings is decreased and it may be considered under this head of restricted 
sexual selection. Evolution is produced if among the mated there is an 
inheritable difference between those of early or normal mating time and 
those of retarded mating time. Such evolution is of course in the direction 
away from the attributes of the retarded unless counteracted by lethal or 
fecundal selection. 

The late married certainly do differ from other mates and the applied 
eugenist is concerned because one important group of the late married 
result from prolonged full time education, especially in the medical profes¬ 
sion. We may therefore well plead for (a) more part time education in pro- 


MATE SELECTION 


425 


portion to the full time education in those of marriageable age, (b) a 
restriction of the requirement of full time medical students to two years of 
college before their five years of medical study, (c) laws prohibiting marriage 
under eighteen to help correct the balance from the other end. 

The leading conclusion of this paper as it affects applied eugenics is “ Se¬ 
lect the best available as to the most important attributes. ,, To this older 
well known maxim we now must add a second “ All superiors should mate.” 


DISCUSSIONS OF PAPERS READ AT THE CONGRESS 


SEX DETERMINATION IN ROTIFERS 
A. F. Shull 

George H. Shull: I would like to suggest that there is no necessary inconsistency 
between the cases so often demonstrated in the rotifers, in which sex has appeared to rest 
upon some condition of the environment, as nutrition for example, and these new cases 
in which it seems to be traceable to the events which take place at the period of meiosis; 
for if we assume that in all these cases the proximal cause is an environmental (nutri¬ 
tional?) one, it is only necessary to recognize that the chromosome constitution is one of 
the necessary elements entering into the production of various enzymes or other chemical 
substances and that difference of constitution between two members of one pair of chro¬ 
mosomes may result in just the difference in enzymes or other chemical substances, which 
is effective in determining the development of the one sex instead of the other. 

It should also be noted that the failure to discover a difference in chromosome number 
in the two sexes is not proof that the difference between the sexes is not based on chromo¬ 
some differences, for in many other species there is no differences in number of chromosomes 
in the two sexes, since the one sex carries an XX pair while the other has a corresponding 
XY pair. 


CONTROL OF SEX IN CLADOCERA 
A. M. Banta and L. A. Brown 

A. F. Shull: It may be possible to explain the relatively smaller effect of greater 
crowding than of less crowding by work on rotifers which shows similar results. The con¬ 
centration of the excretory products would be proportional to the number of individuals. 
Now, in rotifers, in which the effects of a long list of chemical substances upon the life 
cycle have been tested, it was several times shown that a strong solution of a given sub¬ 
stance produced a smaller visible effect than did a weak solution of the same substance. 
It would not be surprising, therefore, to find that great crowding of Cladocera modifies 
the life cycle in some respects less than less crowding does. 

Major Darwin: Has the direct bearing of this subject on eugenics been considered? 
Guinea pigs and rats under crowded conditions breed freely, and under uncrowded condi¬ 
tions do not breed so well. Crowding can affect sexual reproduction. This material may 
have a bearing on eugenics, as an explanation of the effect of crowding in slums. 

Dr. Banta: I have not considered the eugenical bearing. This might not apply. 
Certain specific cultural media also lead to the production of males. 

426 


DISCUSSIONS 


427 


E. G. Conklin: Many rotifers living under conditions which, humanly speaking, 
would be considered bad, reproduce freely. Those living under good conditions sometimes 
produce no males. Phyllodena roseola have been kept under the best conditions and have 
gradually ceased reproduction and the females become entirely sterile. Occasionally 
the water dried up and they underwent encystment, and when they were soaked again, 
the females immediately began to reproduce actively. This might have eugenical bearing 
considering what we in America are undergoing now. 

LINKAGE WITH LETHAL FACTORS, THE SOLUTION OF THE OENOTHERA 

PROBLEM 

George H. Shull 

R. Ruggles Gates: I would like to ask what was the source of rubricalyx used. 
I have used rubricalyx in my experiments and have gotten an entirely different set of 
results. 

Dr. Shull: I made the crosses discussed in this paper with material from Sutton and 
Sons, Reading, England, to whom Dr. Gates sold his purebred stock of this species. The 
source of material is wholly immaterial, however, in experiments such as I have been con¬ 
ducting in the Oenotheras for a number of years, for I have been dealing with rubricalyx , 
Primarily as a unit factor, only incidentally as a taxonomic unit. 

Dr. Gates: In my experiments I used the original rubricalyx. The rubricalyx which 
I sold to Sutton has not remained pure. I have had occasion more recently to test 
Sutton’s material of this species, and I found that it showed evidence of crossing with some 
other species. In 1914 I published a complete account of the F 2 and F8 generation, so 
I am familiar with the behavior of this species and able to identify hybrid conditions. 

INHERITANCE OF MENTAL DISORDERS 
A. J. Rosanoff 

Stewart Paton: There are a great many difficulties which the alienist has to meet 
in dealing with the problems of heredity, and I was greatly pleased in Dr. Rosanofif’s 
paper that he was so conservative in drawing definite conclusions. Mental disorders are 
much more complicated than the casual observer believes them to be. 

A mental disorder is a more complex disorder than a maladjustment at the psycho¬ 
logical level. It is an imperfect adjustment of the whole organism. The average lay 
person believes that insanity is a disease of the brain, but it is a disease of the entire 
organism. We have been so accustomed in recent years to follow Hughlings Jackson and 
speak of levels of activity, that we have begun to think of different levels of insanity. 
Insanity is the way that the entire organism reacts to a difficult situation. The final test 
of sanity or insanity is what the person does. 


428 


DISCUSSIONS 


THE GENETIC SIGNIFICANCE OF THE ALCOHOLIC TREATMENT OF 

WHITE RATS 

E. C. MacDowell 

A SUMMARY OF THE EFFECTS OF EXCESSIVE ALCOHOL TREATMENT 
ON HEREDITY AND DEVELOPMENT IN MAMMALS 

C. R. Stockard 

E. M. East: It seems to me that another point might be brought up. These experi¬ 
ments are, in general, made with alcohol because it has been known for a long time that 
excessive alcoholism does effect the organism. We could use other materials and get 
effects. To any one with a knowledge of chemistry it is not necessary to conclude that 
anything like the same effects could be found with the use of the same quantities of 
another material. It is worth while noting that the use of a small quantity might have 
one effect and that of a larger quantity a different effect and a conclusion ought not to be 
drawn, as Dr. Stockard has said, by the sociological press as to the effects of a similar use 
of alcohol by human beings. 

J . Arthur Harris :There is a possibility of an advance in the analysis. Dr. Stockard has 
called attention to a real difference in the death rate in the two groups of animals. There 
should be associated with that a very great difference in the variability of the offspring. 
Has Dr. Stockard made a study of the offspring in the cases where the mortality is high or 
especially low? 

Dr. Stockard: There is not a great variability. 

G. H. Shull: It is not quite clear to me that Dr. Harris’ point is well taken at this time. 
It seems to me that if there is a considerable variation due to this condition under which 
the organisms have developed there might be increased instead of decreased death rate. 
It seems to refer any variability back to internal constitution and leaves nothing for the 
environment that has been introduced into the experiment. In some of them there are 
more favorable conditions enjoyed than there would be under normal conditions and 
such conditions would serve to decrease rather than to increase the death rate. 

Lucien Howe: I want to give a word of appreciation for the work of these men. 

Raymond Pearl (chairman): Is there further discussion? If I might be permitted 
from the chair I should like to say just a word on this myself, because some years ago 
I did some work on alcohol, and I have been very much interested in the work which 
Dr. Stockard is doing. I think that any difference between Dr. Stockard and myself 
that there has been is a difference in emphasis. Alcohol unquestionably has a selective 
action upon germ cells. I got, finally, a race of chickens that any poultryman would 
call a race highly superior to normal chickens. Those that were defective were eliminated, 
and so far as the race was concerned, have no consequences for the future. I wish to 
make it clear that I am not criticizing Dr. Stockard’s results. I have the greatest admira¬ 
tion for the intelligence and scientific acumen with which he has done his work. I 
am really trying to emphasize another point of view which just as much deserves emphasis. 

I thank you for letting me enter the discussion from the chair. 


DISCUSSIONS 


429 


Dr. MacDowell: Why was not the selective effect of alcohol immediately apparent 
in the guinea pigs, if this action of the alcohol accounts for the condition of the fourth 
generation? In the experiments with rats the number of litters produced by the rats 
actually treated was 60 per cent smaller than that produced by the brothers and sisters 
in the same period, while the untreated descendants of the treated rats produced in 
the next generation from 30 to 50 per cent more litters than the controls. Another point 
that the rats bring out very clearly is that controls in one generation do not serve as controls 
in another generation. I believe personally so far as this point is concerned that the 
results that Stockard’s work bring out are very valuable but this is a point that must be 
brought to mind. 

Dr. Pearl (chairman): Before we disperse, I should like to call your attention to the 
fact that there are other Sections in operation in the West Assembly Room in the Hall of 
the Age of Man. 

I should like to say just one thing, if I may, in closing. I think it may be proper for 
me to say it, since the present Chairman had nothing whatever to do with the arrange¬ 
ment of the program or with the planning of the program. I think all Geneticists will 
agree, and all that I have talked with have agreed, that we have had a very remarkable 
program in Genetics here, and that it has marked many advances in our knowledge ©f 
Genetics, and that it gives the impression that Genetics is moving rapidly in this country. 

I am sure that I voice the sentiment of all interested in Genetics, and all those who 
have been members of the audience here when I say that we thank the Secretary of the 
Section of Human and Comparative Heredity, Dr. Helen Dean King, for arranging such 
a wonderful program for us. 

INTERMARRIAGE OF BLOOD RELATIVES IN THREE OLD NEW ENGLAND 

COMMUNITIES 

Ruth Moxcey Martin 

R. H. Johnson: There is an old saying “Blood will tell.” The difficulty is to say what 
blood will tell. Although there were these few families, doubtless if there were discrim¬ 
ination against all cousin marriages, the town and family would have given us a better 
future. 

Dr. Davenport: There is no question which is more commonly asked of the Eugenics 
Record Office than that concerning cousin marriages. We receive a good many inquiries 
about proposed matings, and I presume that in over half of these cases, these inquiries 
have been made because of a proposed cousin marriage, and the inquiry has been urged 
by the families of the young people because such a marriage has in the past proved unfor¬ 
tunate in the family. We always respond that it is impossible to lay down any general 
rule, that it is necessary to know the family connections, and especially the traits of the 
common blood. We can only point out instances like those which have been so well 
described by the last speaker, where the presence of degeneration or obvious defect of 
the common blood is sure to reappear in the offspring of a cousin marriage. But if, on 
the other hand, there is no obvious defect in recent generations in the close cousins of the 
blood considered, then we have no reason to expect defects in the offspring. On the 


430 


DISCUSSIONS 


contrary the mating may be regarded as a eugenic one, and we have occasion to call 
attention to instances of cousin marriage which have produced only uniformly good 
progeny. We have often used the example of Charles Darwin who married his first cousin 
and who produced such a family of illustrious sons and daughters. 

THE LEARNED BLACKSMITH 
H. J. Banker 

Wilhelmine Key: I want to say just a word. I think we are peculiarly indebted to 
Dr. Banker for this exceedingly careful and detailed study that he has made and for 
the cautious conclusions to which he has been led, because I think genealogists as a whole are 
frequently criticized for leaping at conclusions, and we have in Dr. Banker’s paper an 
example by which we may all profit. 

In this connection, I should like to say too, that any student of genealogy findsabundant 
examples of the type that Dr. Banker has brought before you this morning, that is, a con¬ 
centration in a given personality of gifts which characterized many of the more obscure 
members of the network to which he belongs. Frequently too, we meet in the biographies 
of our eminent men a disinclination to ascribe to ancestral influence abilities which might 
otherwise be regarded as the product of their own striving. 

As an example of this we may cite the late Theodore Roosevelt. He always said that 
he came from good fighting stock, but beyond this, he was inclined to give scant weight 
to heredity as a factor in his course of development. However, if you go back to Theodore 
Roosevelt’s ancestors, you find abundant occurrence of the traits that made for his 
excellences, to name only a few, we find a love of nature and a love of truth, fearlessness 
and a dauntlessness that made them stand for what was right and worth while, and lit¬ 
erary ability of a high order. Not only do we find such traits as these, but we find on, 
both the maternal and the paternal side their combination in a personality that might 
stand as a prototype of Roosevelt himself. 

On the maternal side there was Archibald Bulloch, who was his great grandfather. He 
was governor of Georgia in Revolutionary times and espoused the patriot cause when 
scarcely any of his associates did, pursuing a course that was notable for its courage and 
independence. 

On the paternal side we find Robert Barnwell Roosevelt who was his father’s brother. 
He was a New Yorker, established the New York Fish Commission and for twenty years 
served as Fish Commissioner without pay. He wrote books on the fishes and game birds 
of North America. You know how strong those tastes were in Theodore Roosevelt. 
Fie was, moreover, an ardent reformer, and edited a paper which was opposed to the Tweed 
ring; elected to Congress, refused to obey his party’s behest. In his leisure hours, he wrote 
poetry and stories that show a decided mellowness and play of human feeling. I think 
he had a mellowness and geniality which you did not quite get in his nephew. 

Roosevelt’s remarkable development appears to have come through the possession 
of these salient temperamental and intellectual traits associated with a physiologic condi¬ 
tion where everything he undertook had to be done with all his might and main. This 
condition was also characteristic of other members of his family. Then Roosevelt came 
into public life when his dominant traits of fearlessness and rebellion against domination 
by a party were becoming a large part of national feeling. We get here an action and 
reaction between individual tendency and social and economic conditions, with success 
and applause which serve to encourage and intensify the individual tendency. This 
constitutes leadership in a critical time in national history. 


DISCUSSIONS 


431 


SOME FAMILIES AS FACTORS IN ANTI-SOCIAL CONDITIONS 
Amos W. Butler 

Edward Wallace Lee: This paper is a most valuable contribution. Feeble-minded¬ 
ness, so far as I can observe, is increasing. When we recognize this condition and recognize 
that it is increasing, what is the remedy for us? 

This is the great problem that concerns all who are interested in these conditions. 
For some thirty years I have been interested in this subject and have had an opportunity 
to study these cases and where indicated have resorted to castrating to prevent the 
propagation of this class of individuals, and those that I have treated were thus prevented 
fronrbecoming a menace to the community. 

Another factor in this condition that is not receiving the attention it should, especially 
from the medical profession, is, that the physical condition of the feeble-minded is not 
given sufficient attention. There may be a condition of the mind that is not dependent 
upon the body, but I doubt it, at least, I cannot imagine it. I cannot imagine a defective 
mind unless there is a physical defect existing somewhere either in the brain itself or in 
some other portion of the body. I know that a diseased or mal-functioning intestinal 
tract will produce a condition which might be called feeble-mindedness at least it disturbs 
the mentality. The thyroids and other ductless glands when not properly functioning 
may produce the same deleterious effect on the mind. 

These feeble-minded people are not given the physical attention that their condition 
demands. Criminals are not given the physical attention that their condition demands. 
They may be considered and be pronounced feeble-minded; their minds may have been 
examined; psychological examinations may have been made, but their bodies have not been 
given the attention that they should honestly and scientifically receive. 

THE TRIBE OF ISHMAEL—A STUDY IN CACOGENICS 
Arthur Estabrook 

Dr. Paton: Do the symptoms characteristic of the maniac depressive type of per¬ 
sonality or the schizophrenic type seem to be responsible for the behavior of any of 
the cases that you have reported? 

Dr. Estabrook: There has been practically no insanity in the Ishmael family. The 
little that has appeared has been largely due to syphilis. The characteristic mental 
status of the Tribe has been that of a high grade feeble-mindedness. 

Dr. Paton: I should like to ask whether there was any detailed physical examination. 
Might it not be possible to correlate some of the results of the physical examination with 
their mental traits? 

In the aviator one of the signs of physical unfitness for work was loss of interest in the 
work. Dr. Mac Lake in handling four thousand cases among the aviators found that the 
loss of interest in the work was often closely correlated with the change of the dermato- 

graphic reaction on the skin. f 

Possibly these vagrant types of personality might show interesting correlations oi 

physical and mental symptoms. 


432 


DISCUSSIONS 


Dr. Estabrook: I should say in answer to Dr. Paton that this type of vagrant person¬ 
ality has been observed and studied physically in but very few cases. From the stand¬ 
point of physical examinations, it has been possible to get clear, good, physical examina¬ 
tion of but few persons, those that have been in institutions, or those who for some reason 
or other it has been possible to bring to some sort of a clinic. This study has not gone 
iDto the finer discriminations because of the magnitude of the problem. 

The Ishmaels are found in ten states and number about fifteen thousand individuals, 
and it has been possible to get only the roughest data concerning the more or less pro¬ 
nounced traits. 

Franklin H. Giddings: Has anything been done to prevent the increase of the tribe? 

Dr. Estabrook: From the standpoint of the Record Office, this study has been purely 
academic. From the standpoint of what can be done to prevent increase, I might say 
that some data from the Record Office have been placed in the hands of the Board of 
State Charities in Indiana and other states; it has been given to charity organization 
societies, to different state institutions, and these other workers have done what they 
can to take care of these feeble-minded situations and to see that those that can be placed 
in institutions are so placed. From the standpoint of prevention of increase, I think 
the present problem is the placing of the information before the public, thus enabling them 
to take care of these defectives properly and with an intelligent understanding of the 
problems they are attempting to solve. 

It is not the function of the Record Office to carry on any definite propaganda. Its 
function is purely academic. 

Dr. Lee: Did you report on the whole tribe or just select individuals of that tribe? 
I was wondering if there are any good Ishmaels. 

Dr. Estabrook: In the last decade or two there are a few good Ishmaelites. I think, 
of the whole group, approximately two or three per cent might be classed as average 
persons, socially and intellectually, and when I use the term average, I am using it accord¬ 
ing to the standard worked out by the United States Army in its classification of draft 
quotas. I can think of but two or three individuals who have become what we would 
call exceptionally good individuals, socially adequate, from among the Ishmaels. 

About 90 per cent of the total group, and even more, are what we would call socially 
inadequate. The Ishmaels have been more or less under the public eye to a certain 
extent for about forty years, especially in the cities, and we do not find many among 
them who reached the so-called average class. 

Prof. Giddings: Can you tell us in a word how it became possible to get a piece of 
scientific work of that character and extent done in state political institutions of today? 

Dr. Estabrook: My association with state institutions and state boards has been 
entirely as a representative of the Record Office, and in going to these various places 
I have been received most cordially. At the Board of State Charities in Indiana, Mr. 
A. W. Butler, Secretary has cooperated very extensively. 

I might say too, that in working in the various states of the Union, and these include 


DISCUSSIONS 


433 


over half of the states, I have been received most cordially by those persons to whom I 
have gone, and information has been very readily and quickly given. 

At the time I was working on the Jukes, the State Commissioner of Prisons and the 
State Board of Charities in this state (New York) turned their office material over, 
even the Bertillion measurements of criminals in this state. 

Political conditions have had no influence, lliat has been due to the fact, I think, 
that I am associated with a private organization. 



INDEX 


Ability, intellectual, 346 
native, 378 

Acrididae, North American, 66,67,68,69,73 
Adams, John, 331 
Adams, John Quincy, 331 
Agreement, faculty of, 378 
Alden, John, 330 
Alethal forms, inferiority of, 87 
Allotment, 264 
Ancestor, 307, 309 
Anderson, W. S., 297 
Animals, adaptation to environment, 40 
cavemicolous, 45 
Anticipation, 221 
Aptitudes, 408 
Arcella, 62 
Arches, 199 
Aristogenic type, 340 
Arms, of Izquierdo family, 352 
Arteriosclerosis, 170 

Bagg, Halsey J., 150 
Banker, Howard J., 306, 340 
Banta, Arthur M., 142 
Bartlett, J. Gardner, 319 
Beauty, 419 
Beggars, 401 

Bell, Alexander Graham, 21 
Belling, John, 84 
Biologists, 54 

Birth rate, in France, 243,250 
factors determining, 252,256 
Birth weight, 268 
Blakeslee, A. F., 82 
Blindness, 193 
Bonnevie, Kristine, 198 
Bradford, William, 330 
Brain tumor, 170 
Brain, organic disease of, 222 
Bridges, Calvin B., 76 
Broodiness, 212 


Brown, L. A., 142 
Burritt, Elihu, 340 
Butler, Amos W., 387 

Cady, Lee D., 174 
Cancer, inheritance of, in man, 186 
in mice, 182 
tendency to, 184 
Capacity, motor, 233 
sensory, 233 
Celibacy, 422 
Cells, germinal, 35 
Ceriodaphnia, 145 
Characters, acquired, 30,62,63 
fixed and unfixed, 129 
Chorea, 297 

Chromosomal condition and body charac¬ 
ter, correlation of, 69 
Chromosome complex, evolution of, 65 
Chromosome groups of mutating and 
hybrid species, 72 

Chromosome materials, aberrations in, 76 
Chromosomes, 82 
attraction between, 84 
in Oenothera, 86 
variation in numbers of, 68 
Cladocera, 142,143,145,149 
Coaptation, 45 
Coleoptera, 48 

Conception, prevention of, 424 
Conification, 312, 314 
Consanguinity, 276, 289 
Cornea, 191 
Cotton, Henry A., 170 
Crime, hereditary factors in, 413 
Crossing, 267 

Crowding, effect of, on reproduction, 144 
Cuenot, L., 29 
Cytoplasm, 29, 84 

Danforth, C. H., 120 
Daphnia, 142, 143, 144, 145, 146 


436 


INDEX 


Darwin, Leonard, 5 

Darwinian evolution by mutations, 115 
Datura, 79, 82 

Davenport, C. B., 20,170,171 
Death rates, 243 
at Oneida Community, 385 
Degeneracy, 282 
Degeneration, 271 
Dementia praecox, 223, 226 
Difflugia, 60, 61 
Diploid plants, 82 
Dominants, 79,134, 213 
Drosophila, 76, 88, 108 
bar-eye in, 113 

Egg, of domestic hen, 212 
of fish, 43 

schematized fertilized, 130 
Endocrine glands, relation to develop¬ 
ment, 25 
Endogamy, 286 
Energy, 340,343,346 
Environment, 245 
Epilepsy, 223, 226 
inheritance of, 24 
Estabrook, Arthur H., 398 
Eugenical field workers, 21 
Eugenical records, 22 

Eugenical societies, aims and methods of, 
5, 16 

educational work of, 10 
research work of, 8 
Eugenics, 244, 247 
attitude toward pauperism, 396 
research in, 20 

Eugenics Record Office, 20, 170 
Evolution, 30 
theory of, 41 
Ewing, James, 166 
Exogamy, 285 
Experimentation, 236 
Eye defects, 163,191 

Families, degenerate, 389 
Family history, 306 
Family tree, 349 
Fecundity, 26, 248 
in domestic hen, 212, 216 
low, 410 


Feeble-mindedness, 298 
inheritance of, 23 
Fertility, 216 

effect of social situation on, 262 
in inbred strains, 273 
Finger prints, 198 
Fish, 42 

Fisher, R. A., 115,195 

Galton, Francis, 21,198 
Gamete lethals, demonstration of, 96 
Gametes, 88 
Garfield, James A., 332 
Gates, R. Ruggles, 100 
Gene, 78 
theory of, 120 
Genealogy, 306 
of Elihu Burritt, 341-346 
of Izquierdo family, 348 
Genetic differences, analysis of, 102 
Genetics, 7 
human, 21 

Genetics and adaptation, 29 
Gentry, 316, 321 
Germ plasm, 276, 297 
Grant, U. S., 332 
Greene, Elizabeth, 413 
Groups, social, 312, 314 
Gudernatsch, J. F., 151 
Guinea-pigs, inbreeding in, 180, 266 
Guyer, Michael F., 192 
Gypsying, 401 

Hadrobracon brevicornis, 102 
Haploid parthenogenesis, 102 
Hardiness, 376 
Harriman, Mrs. E. H., 20 
Hens, 34 

Hereditary mechanism, 129 
Hereditary traits, in man, 120 
Heredity, 135,138, 245 
relation to pauperism, 392 
relation to tuberculosis, 178 
Heterozygous types, 116,196 
Homozygous types, 116, 196 
H<pwe, Lucien, 191 
Hurst, C.C., 212 


INDEX 


437 


Identical twins, 195 
Imagination, 233 
Impregnation, artificial, 421 
Inbreeding, 270, 273, 285 
in guinea-pigs, 266 
Industries, 259 

Inheritance, of cancer, 182,186 
of eye defects, 191 
of mental disorders, 226 
of musical traits, 231,239 
laws of, 7,9 

Insanity (see Mental disorders) 
Institutions, inmates of, 388 
Intensity-discrimination, 240 
Intermarriage, 278 
Ishmael, Tribe of, 398 
Ishmaelites, 398 
Isolation, 297 
Izquierdo, J. Joquin, 348 
Izquierdo family, origin of, 350 

Jennings, H. S., 59 
Jimson weed, 82 
Johnson, Roswell H., 416 

Karyokinesis, 135 
Key, Wilhelmine E., 405 
Kimball, Sarah Louise, 329 
King, Helen Dean, 270 

Lamarck, 41 

Lamarckian examples, 38 
Legislation, eugenical, 15 
Legrand, Louis, 129 
Lens, crystalline, 34 
Lethal factors, linkage with, 86 
Lewis, Paul A., 178 
Licentiousness of Ishmaelites, 401 
Lidbetter, E. L., 391 
Life cycle, changes in, 138 
Lineage, Spanish, 351 
Mexican, 358 

Linkage, Oenothera, 89, 91, 92 
Litter, 303 
Little, C. C., 186 
Loeb, Leo, 182 

Longevity, 377 , 


Longfellow, Henry W., 336 
Loops, radial, 200 
ulnar, 200 

MacDowell, E. Carleton, 168 
Malthus, 253 

Manic-depressive psychoses, 224, 228 
March, Lucien, 243 
Marriage, types of, 287 
among Indians, 294 
early, 411 

Marriage rate, of insane, 220 
Martin, Ruth Moxcey, 278 
Mate selection, 416 
preference in, 418 
Matings, assortative, 408 
cousin, 28 

Maturation, of egg, 140 
Mayflower, 329 
McClung, C. E., 65 
McCulloch, Oscar, 21, 399 
Memory, 233 
tonal, 240 

Mendelian factors, 212 
Mendelian traits, inheritance of, 22 
Mendelism, 40,45, 59,115, 275 
Mental defectives, 388 
Mental deficiency, 226 
Mental disease, 218 
transmission of, 223 
Mental disorders, 218, 226 
inheritance of, 170 
Metaphase, 84 
Meyerson, A., 218 
Mice, 303 
Moina, 142 
affinis, 143, 145 
macrocopa, 143,144,145,147 
rectirostris, 143, 145 
Muller, H.J., 106 

Musical talent, measurements of, 231, 234, 
240 

prediction of, 238 
Musicians, 240 
Mutation, 29, 76, 81,106 
in man, 120 
in Oenothera, 98 


438 


INDEX 




Nanella stature, 92 
Nativity, of runaway girls, 414 
Neoplasmic growths, 187 
Neuropathic heredity, 219 
New England, old families of, 315 
Noyes, George W., 374 
Noyes, Hilda H., 374 
Noyes, John H., 374 

Oenothera, 79,86,98,100,107 
Oenothera problem, solution of, 86 
Officer-yeoman, 316,322 
Offspring, endowments of successive, 303 
Oneida Community, 374 
Osborn, Henry Fairfield, 1 
Ovary, secretion of, 183 
Overexpansion, 309 

Palpation, 174 
Papillary ridges, 198 
Paramecium, 60 
Paranoia, 223, 226 
Paraplegia, 158 
Parthenogenesis, 139,142 
Pauper stocks, pedigrees of, 391 
Pauperism, 391 
in Indianapolis, 399 
Paupers, 282 
chronic, 394 
occasional, 394 
Pedigree chart, 310 
Pilgrims, 329 
Pioneers, 278 
Pitch discrimination, 239 
Polydactyly, 121 

Population, of France, 243, 256, 263 
Pre-adaptation, 42,44 
Predisposition to cancer, 186 
Professions, 258 

Progeny of cancerous individuals, 188 
Prostitution, 402 
Psychoneuroses, 226 

Rabaud, Etienne, 303 
Rabbits, 34, 193 

Radium, effect of, on mammalian develop¬ 
ment, 150,151 


Rats, albino, 270 
alcoholic treatment of white, 168 
Recessives, 35,213 
Reproduction, mode of, 140 
Repute, 420 
Reverse mutations, 113 
Root, Elihu, 333 
Rosanoff, Aaron J., 226 
Rufer family, 406 
Runaways, adolescent, 413 

Seashore, Carl E., 231, 239 
Selection, artificial, 247 
natural, 57,115,246 
sexual, 416 
Self-adaptation, 408 
Sex determination in rotifers, 138 
Sex-linked traits, 23 
Sex phenomena in Cladocera, 142 
Sexual dominance, 136 
Shull, A. Franklin, 138 
Shull, George H., 86 

Simocephalus exspinosus, 143,144,145,146 
serrulatus, 143,145 
vetulus, 143,145 
Size-inheritance, 100 
Social control of heritable factors, 405 
Spermatogenesis, 161 
Spermatozoa, 161 
Spinden, Herbert J., 285 
Standish, Miles, 330 
Stanton, Hazel M., 239 
Statistics, 186 
Sterility, 423 
Sterilization, 14 
Stirpiculture, 374,386 
Suicide, 278 

Supracondyloid process, 174 
incidence of, among insane, 175 
Surnames, 281 
Syndactyly, 121 
Synthesis, autocatalytic, 77 
Syphilis, 170, 192 

Taft, William Howard, 334 
Terry, R. J., 174 
Tetraploids, 82 


i 


INDEX 


439 


Time discrimination, 240 
Toxemia, cerebral, 172 
Transmission of character, 33 
Triploids, 82 

Tuberculosis, 178,244,265 
infectious nature of, 178 
Tumorrate, 182 
Twins, 26, 61 
genesis of, 195 

Unicellular organisms, inheritance in, 59 
Unity, in family history, 308 

Variability, new type of, in plants, 100 
Vigor, effects of inbreeding on, 266 
decline in, 271 


Vivacity, 419 

Webster, Daniel, 334 
Whiting, P. W., 102 
Whorls, 199 

Woods, Frederick Adams, 312 
Wright, Sewall, 266 

X-ray, 150,151 

Yeomanry, 316, 324 

Zeleny, Charles, 113 

Zygote lethals of Oenothera Lamarckiana, 
95 









PLATES 












Plate 1. The Chromosomes of Man 


This chart gives the essential results of a study on human spermatogenesis, made by 
Professor Theophilus S. Painter, Department of Zoology, University of Texas, Austin, 
Texas. Figures 1 and 2 show that there are 48 chromosomes (24 pairs) in the germ cells 
(spermatogonia) of a white man, this number including the body labeled “Y.” The 
negro (figs. 3 and 4) shows the same number of chromosomes and the presence of 
the Y-chromosome. In figures 5 and 6 the chromosomes of the white man and the 
negro are compared. They are alike in general form and in number. Figure 7 shows the 
‘‘reduced” chromosome number of man to be 24. Figure 8 shows the sex-chromosomes 
of man which are of the X-Y type. When such a cell divides, the X-chromosome goes to 
one pole and the Y-chromosome to the other. This is shown in figure 9, taken from a white 
man, and in figure 10 which is from negro material. As a result of this, one-half of the 
sperm will carry an X-chromosome, and one-half will carry a Y-chromosome. Sex deter¬ 
mination in man then is simply a matter of which sort of sperm fertilizes the egg. If the 
sperm carries an X-chromosome, then the resulting offspring is a female, but if the sperm 
carries a Y-chromosome, a son will result. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 1 



THE CHROMOSOMES OF MAN 


WHITE 






■4EGRO JSSSTl 1 


-5BW 


WHi i 




NEGR 


Jt)rtl))3Jtl})VO{(|illlllU((l(t CI (€<<(•(*< (d|i 

6 


- 1 ' 4 $^ 














Plate 2. The Catlin Mark 

Inheritance of an unusual opening in the parietal bones. By Dr. William M. 
Goldsmith, Professor of Biology, Southwestern College, Winfield, Kansas. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 2 




rO 


03 


S-o- 


Or 


*4 




EL 


«ww 





































































Plate 3. Inheritance of Order of Succession in Development of the Carpal Bones 

Charts showing, for children of each of three families, the order of succession of develop¬ 
ment of the carpal bones, of the wrist. The X-ray photographs were furnished by Dr. 
Prior of Lexington, Ky. The outline diagrams show the order of development of the bones 
in each individual. Eugenics Record Office. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 3 



» 


























Plate 4. Heredity of Longevity 

Stereograms showing relation between age of father at death, age of mother at death, 
and longevity of offspring. 

Upper series , left to right: 

-20. Persons who died under twenty years. “The stereogram relates to 417 persons 
who died under twenty years of age. The figures show the percentage having fathers and 
mothers who died at ages specified. 8.9 per cent had parents who lived to be over eighty 
years of age.” 

20-40. Persons who died twenty to forty years of age. “The stereogram relates to 
354 persons who died twenty to forty years of age. The figures show the percentage 
having fathers and mothers, who died at the ages specified. 4.8 per cent had parents who 
lived to be over eighty years of age.” 

40-60. “The stereogram relates to 351 persons who died forty to sixty years of age. 
The figures show the percentage having fathers and mothers who died at the ages speci¬ 
fied. 8.1 per cent had parents who lived to be over eighty years of age.” 

Lower series: 

60-80. “The stereogram relates to 333 persons who died sixty to eighty years of age. 
The figures show the percentage having fathers and mothers who died at ages specified. 
18.0 per cent had parents who lived to be over eighty years of age. ” 

80-100. “ The stereogram relates to 138 persons who died eighty to one hundred years 
of age. The figures show the percentage having fathers and mothers who died at the ages 
specified. 27.5 per cent had parents who lived to be over eighty years of age. ” 

Alexander Graham Bell. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 4 






















Plate 5. Pedigrees oe Dramatic and Musical Talent 

(1) Dramatic ability—Kemble family. (2) Musical talent—Bach family. 
Prepared by the Eugenics Education Society, London, England. 


EXCEPTIONAL. ABILITY 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 5 


C® 


JrJ\ <>g .-O';: ;V „o- 

o* > 41 <* 

/Jr <2 

O 5 ! ; °V rO^ 

Lc s 


k? 

io^ 


T^w®! 


r<> 3 


-O’ 


. , _ /§-* 
O" -O" '-ST- ^ k * 5 ^ 

-o- ; . -- 0 s o\r°"’ 

Oj ; -CH 

i^kroVr© 1 

®i rO»H cw>- 

♦' c* -CFKV 


0=1 




fOS* o 

fO^r KT- 

I 0? L-t^ 1 " r O ? 
•0‘ J 


-O’ 


O • 



© ~ 

I I 

»- » 


oi 


cc 


l® ® 





















































Plate 6. Heredity of Musical Ability 

These charts, prepared by Dr. Hazel M. Stanton, Eastman School of Music, Rochester, 
N. Y., are representative of the results obtained in an investigation in the inheritance of 
specific musical capacities, which covered six family groups, in which one member of 
each group was known to be conspicuously talented in music. This investigation, initiated 
in the year 1920, is the beginning of the first research in heredity of talent based on quanti¬ 
tative measurements. Four of the Seashore Measures of Musical Talent, the sense of 
pitch, the sense of intensity, the senses of time and tonal memory were given individually 
to members of each family. These measurements were supplemented by qualitative 
information regarding individual case histories and musical experiences, the latter including 
musical environment during youth, musical training and education, musical activity, 
musical interests, and musical memory and imagination. 

On each pedigree talent chart one mating and offspring are presented showing the results 
obtained in the musical measurements, also the ratings assigned for muscial experiences. 
The results of each of the four measurements are expressed graphically in terms of per¬ 
centile rank ranging from 0 to 100. A rank of 98 to 100 is very superior, 90 to 97 is 
superior, 70 to 89 is excellent, 40 to 59 is average, 10 to 29 is poor. The sense of pitch is 
shown in the upper horizontal section of each individual chart, the sense of intensity in 
the second section, the sense of time in the third section, tonal memory in the lower 
section. The ratings of musical experiences are stated in terms of the letter A, high 
rating, the letter C, middle rating, and the letter E, low rating. At the side of each chart 
a brief description is given of the musical expression evinced by each individual charted. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 6 























































Plate 7. Heredity of Harelip and Cleftpalate 


Pedigree charts. 

Upper: Three families showing harelip inherited without cleftpalate. 

Lower: Three families showing harelip and cleftpalate, often both defects 
individual. Eugenics Record Office. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 7 


\ 






















































Plate 8. Inheritance in Epilepsy 




SLx charts illustrating inheritance of epilepsy. In all charts: A, alcoholic; E, epileptic; 
F, feebleminded; I, insane; Ne, neurotic; S, syphilitic; Sx, sex offender; W, nomadic; 
squares, males; circles, females. Black symbols, defective individuals. 

Dr. David F. Weeks, Skillman (New Jersey) Village for Epileptics. 



EUGENICS, GENETICS AND THE FAMILY 


PLATE 8 


SYPHILIS AND E 

SH-O 


©E ,Ai S’: 


0 




<D 


Ha* an 

Epileptic field 


Hysterical 


d. 


1 


died 
1 n 

Infancy 


IN 4 C.FNFRATION^ 

EPILEPSY 

IN 3 GENERATIONS! 

o - v a 

a~o ~Ze <> ■ 1 

d ”6 6 ft 

id O d </' 

c 5 a a • U ® 9 & 3 / 

•; d it A A 6 ftau o a ilf a a a 

* & 4 5 a ?.• vL a a 

- . '4 

♦ mil 

a a o <* ® * .» a 1 


H 


tmc ocrrcr is. O 

IMTSNaiFtCfi IN THE 




MTttfSIFItt JN THE o . 

GCNCRATION \ 0 . ■ Ci> 

f-fwbVfe-t 

rt*** r 4' r ~ 

* ?4$ / : Kjf I 9 


✓ 


<> a a S&ft 1 

3f> 


*T7j r5<3^ 


C^IEEPSY FECBLE-MJHOCONrsS AND. 5EVUAL ! VMG*Ai.:'* 


•$"5“3 6 66 6 

o ofi3 n a Je 

IT 


- -o *6 jfQ&i' :■ 


l JT" $ *tr 


rTT5T51f%i i W 1 *$> 

to n* do u V n o 

a a jj'o 3~r"- r 

s a 


A r Nt 
i -- 


I *“fT 5 a •' 

*~-jisr & ■ a 

sis? A 


seven epileptics 

DESCENDANT TROH ALCOHOLICS OF THE PERIODIC SPREE TYPE 


°/ 

i 


V'i® J '"t; 4 

> / 


© 




£: 


/ \o 

=- -..*».** 

fo A * 

* l O /o 

*c i 

1 xX 

X' ^ibf?" x 


\ 


/ 


o > 


,*»' 


o' -5, 


4 ^/ : • W 


EPILE PSY AND FEE B LEMINDEDNESS 
A FORCED MARRIAr.P 

^ -o T 


» 


o 




6 


L ,1 ' Sn|Ni: 


OQa 

Ml SO? 

„ BAD BAD 

tlMper. tempo? 



F]pi "fS i SpiT 

@ @ i @ 

4 

























































Plate 9. Inheritance of Specific Iso-agglutinins in the Human Blood 

The blood serum of certain persons will cause the red blood corpuscles of certain others 
to stick together (agglutinate) in clumps. Four human blood groups are recognized (I-IV). 
Their properties are described in the lower right hand chart. 

Prepared by Dr. F. L. Reichert, Johns Hopkins University. 


OH- 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 9 












































Plate 10 . Color Inheritance in Corn 


Purple stalk and leaves; kernels of various colors, also tan stalk, leaves and kernels. 
Seven ears of seven colors of pericarp, ranging from deep purple, crimson, pink, tan, brown, 
yellow and white, all with the royal purple husk, showing constant husk color with varied 
colored kernels. 

Corn bred and exhibited by Harvey J. Sconce, Plant Breeder, Sidell, Illinois. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 10 











Plate 11. Measurement of Physical Traits 


Methods of taking measurements, illustrated on a subject. Reading from left to 
right and from above down: 

1. Vertex height. 2. Tragion. 3. Acromion. 4. Radiale. 5. Stylion. 6. Dactylion. 
7. Suprasternale. 8. Tibiale. 9. Internal malleolus. 10. Anterior ilio-spinale. 11. 
Symphysion. 12. Cervicale. 13. Bicristal breadth. 14. Bitrochaner breadth. 15. 
Iliospinal breadth. 16. Chest breadth. 17. Antero-posterior chest diameter. 18. Sitting 
vertex height. 19. Sitting suprasternal height. 20. Head length. 21. Head breadth. 
22. Tracing skull contours. 23. Measurement over a bathing suit. 

Instruments made by Hermann, Zurich. 

Pictures by Professor H. H. Wilder, Smith College. 

At right, a scale for measuring stature in English and metric systems. 

Iowa Child Welfare Research Station. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 11 






























Plate 12. Measurement of Physical and Mental Traits 

Condensed guide to the Binet tests. Method of head measurement. 
From the Vineland (New Jersey) Training School. 



TO THE BINET TESTS 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 12 















Plate 13. Measurement of Mental Traits 

Mental tests used at Vineland. 

Training School at Vineland, New Jersey. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 13 


















Plate 14. Palm and Sole Prints and Their Inheritance 

Left, above. Various sole prints of European-Americans. 

Left, middle. Palm prints of mother and two sons. Diversity in one family. 

Left, below. Father and son, the latter a complete duplicate of the former. 

Four larger charts on right are prints “interpreted,” i.e., covered with lines indicative 
of the individual conditions. These are of duplicate or “identical” twins,—that is, 
twins that have arisen from a single egg. The general character but not the minutiae 
are the same in both members of a set. 

By Prof. H. H. Wilder, Smith College. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 14 

































Plate 15 . Composite Portraiture 


Composite photographs largely made by the late Henry P. Bowditch of Boston. 

First row: Left upper: 60 Wellesley College students. Left lower: Class of ’87, 
Vassar College. 12 Wends and composite. 12 Portland (Me.) physicians and com¬ 
posite. 12 Saxons and composite. 

Second row : College men from Harvard, Amherst, 449 components. Co-composite, 
Harvard Annex, Smith, etc., 287 components. Harvard Class of 1887, 156 members. 12 
Horse-car drivers. General paresis, 8 components (5 men and 3 women), 11 Mathema¬ 
ticians Amherst Class of 1887, 71 components Williams College, 57 components. 16 
Naturalists. 

Third row: Women’s Medical College, 1887, 38 components. Component three 
members of Bowditch family. 12 Boston Doctors and composite. Mt. Holyoke class 
of 1887, 47 components. Harvard Annex, 1887, 47 components. Smith, 1887, 38 com¬ 
ponents. 

Fourth row: Horse-car conductors, 12 components. Sheffield Scientific School, Class 
of 1887. Cornell, 1887, 65 men, 5 women. 30 Members of the National Academy of 
Science. Melancholia, 8 components. Harvard Faculty, 1887, 38 components. 

Bottom row: Upper left: 12 Portland doctors and composite. Lower left: 12 Horse-car 
drivers, Boston ’88, 12 Saxons and composite, 12 Wends and composite, 12 Saxons and 
composite. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 15 











































I 


Plate 16. Pedigree of John Burroughs 

Family history chart, with three portraits and a life mask, of John Burroughs. Also 
portraits of brothers, sisters, parents and other close relatives. At the right, two photo¬ 
graphs of Burroughs; one of them, his last, taken a few days before his death. _ 

By Harry H. Laughlin, Eugenics Record Office. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 16 
























Plate 17. Pedigree of the Caesars 
From the Exhibit of the Eugenics Education Society, London, England 





PEDIGREE OF THE CAESARS. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 17 



k 

S3 
«5 
© 

S3 

® 4 

4 = 

I .1 

is u 
S3 be 

QJ PC 


3 

X) 

L 

S3 

JD 

O 

C 

© 

JC 

PC 

UD 

.2 

*3 

1 

S 

© 


CO 




fC 5 
> > 


> 

© 

3 


CC 

> 


S3 

c. 

© 

C* 

S 

pC 


S3 

co 

© 

«3 

a> 


£ 

© 

z 


PC 

> 


□- uf-i 



Im 
C3 

?/} 

* 4i 

f* cs 

- 

O. ** 

o = .2 

""3 3 

-5 —I 

3 _ M « 

PC PC PC < 


© 

Cl 

E 

£ 

© 


c 

o 

'u 

c3 

to 

© 

« 

a) 


co 

a 

’> 

S3 

<-> 

o 

O 

to 

J3 •- 
-*->» C5 

ai 


pc 

c 


83 

‘> 

S3 

** 

o 

© 


>> 

C 

O 

-*«> 

c 

PC 

o 

' S3 

s 

© 


«* CD OQ © CO 


C 

© 

£ 

*u' 

£ 

O 

PC 


S3 

to 

© 

e3 

a) 

to 

3 

+* 

Cfl 

3 

-be 

3 

PC 


to 

3 

*3 

3 

_C 

5 £ 
to © 

.2 Z 

t- 

u 

£> 


<a 
> 

J H 

3- - 

PC PC © 

« 


L I *c y 
























Plate 18 . The Old Americans and the Tribe of Ishmael 


Upper: Physical proportions and physiological characteristics of females as compared 
with males, among the old Americans. Top line of figures gives the rates of female to 
male dimension. From Dr. A. Hrdlicka, U. S. National Museum. 

Lower: The tribe of Ishmael, by Dr. A. H. Estabrook. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 18 






















Plate 19. The Jukes 


The Juke charts compare the family as known to Dugdale in 1875 and again to A. H. Esta- 
brook in 1915, forty years later. Dugdale, 1875, while inspecting the county jails of New 
York State, discovered this family of criminals, prostitutes and paupers, studied their 
family history and gathered data concerning seven hundred persons descended from 
“Margaret, called the Mother of Criminals.” In 1915 Estabrook studied the same family 
of people to ascertain the changes in social and mental status which had taken place in the 
intervening forty years. The charts show the two sets of data, one of course inclusive of 
the other, comparing the family at the different periods and showing that the Jukes are 
still a serious burden to the coummunity. A few Jukes have risen from the mire and are 
now socially adequate persons. Pictures of various members of the family and their 
living conditions are shown. By A. H. Estabrook. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 19 



" 1 IT $ * 














Plate 20. The Nams 



The Nams are a set of feeble-minded folk living in the northern part of New York 
state. They are characterized by illegitimacy, prostitution, consanguinity and feeble¬ 
mindedness. They number about two thousand persons, practically none of whom has 
become socially adequate. The majority of the family is still reproducing its own kind of 
dysgenic folk. The charts show pictures of the folk and their homes and general habitat. 
A. H. Estabrook, Eugenics Record Office. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 20 












Plate 21 . Mentality and Delinquency 


Relation of illegitimacy to parental mentality and infant mortality, also of delinquency 
to mentality. Children’s Bureau, U. S. Department of Labor. 


EUGENICS, GENETICS AND THE FAMILY 


PLATE 21 




















Plate 22. The Brains of Criminals 


Photographs of criminal brains, showing great variety of forms. Part of exhibit of 
Massachusetts Department of Mental Diseases, by Dr. Myrtelle M. Canavan. 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 22 


MASSACHUSETTS 


DEPARTMENT 


OF 



576. ALCOHOLIC VAGRANT. 
BRAIN NARROW. SIMPLE. 
WT. 1550. 

MOTHER DIED INSANE. 
CANADIAN. 


530. SEX PERVERT. VAGRANT. 
BRAIN BROAD SHORT ANOMALOUS. 
WT. 1400. 

GOITER. 

PARENTS UNKNOWN. 
AMERICAN. 



MENTAL DISEASES 
EXHIBITS 
PICTURES OF 50 
CRIMINAL BRAINS 

• e Je 

42 


NORMAL ! 

♦ 




954. RAPE. 

BRAIN LONG. UNEVEN. 

WT. 1230. 
PARENTS UNKNOWN. 
AMERICAN. 


585. ALCOHOLIC VAGRANT. 
BRAIN LONG, SIMPLE. 
WT. 1230. 

PARENTS UNKNOWN. 
CANADIAN. 


553. VAGRANT. 


788. PARETIC VAGRANT. 


BRAIN LONG SQUARE ENDED SIMPLE 
WT. 1370. 

PARENTS AND SIBLINGS NORMAL. 
IRISH. 


BRAIN LONG NARROW UNEQUAL.' 
ATROPHIC. WT 1200. 
PARENTS UNKNOWN. 
ITALIAN. 






































Plate 23. Marriage and Birth Rate in Relation to Immigration 

Marriage, fecundity and immigration and their significance for the nation. 
Charts furnished by the Race Betterment Foundation. 


CONCENTRATION ON ISLANDS 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 23 



/ 















































Plate 24. Approaching Extinction of “Mayflower” Descendants 

The declining birth rate and its consequences. (1) Approaching extinction of May¬ 
flower Descendants. (2) Declining birthrate among Mayflower Descendants. 

(3) Varying fecundity of the Brewster family. Eugenics Record Office. 


NUMBER Of DESCENDANTS 


EUGENICS. GENETICS AND THE FAMILY 


PLATE 24 



l 
























































































































































